Russian Entomol. J. 12 (1): 11–58 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2003

Vibrational communication in leafhoppers from Ulopides subfamilies group (Homoptera: Cicadellidae) and Membracidae with notes on classification of higher taxa

Âèáðàöèîííàÿ êîììóíèêàöèÿ öèêàäîê ãðóïïû ïîäñåìåéñòâ Ulopides (Homoptera: Cicadellidae) è Membracidae ñ çàìå÷àíèÿìè ïî ñèñòåìàòèêå âûñøèõ òàêñîíîâ

D.Yu. Tishechkin Ä. Þ. Òèøå÷êèí

Department of Entomology, Faculty of Biology, M.V.Lomonosov Moscow State University, Vorobjevy Gory, Moscow 119899 Russia. E-mail: [email protected] Êàôåäðà ýíòîìîëîãèè Áèîëîãè÷åñêîãî ôàêóëüòåòà Ìîñêîâñêîãî ãîñóäàðñòâåííîãî óíèâåðñèòåòà èìåíè Ì.Â.Ëîìîíîñîâà, Âîðîáüåâû Ãîðû, Ìîñêâà 119899 Ðîññèÿ. E-mail: [email protected]

KEY WORDS. Cicadellidae, Membracidae, Cicadina, Idiocerinae, Macropsinae, Agalliinae, Adelungiinae, Megophthalminae, Ulopinae, leafhoppers, taxonomy, signals, bioacoustics, vibrational communication. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ. Cicadellidae, Membracidae, Cicadina, Idiocerinae, Macropsinae, Agalliinae, Ade- lungiinae, Megophthalminae, Ulopinae, öèêàäêè, òàêñîíîìèÿ, ñèãíàëû, áèîàêóñòèêà, âèáðàöèîííàÿ êîììóíèêàöèÿ.

ABSTRACT. Oscillograms and descriptions of vi- Ossiannilsson [1949]. Nevertheless, until recently there brational signals of 50 species of leafhoppers belong- was very few if any information on vibrational commu- ing to Idiocerinae, Macropsinae, Agalliinae, Adelungi- nication in most part of families, subfamilies and tribes inae, Megophthalminae and Ulopinae (Homoptera, Ci- of this group. Therefore, some time ago I started a cadellidae) and of 3 species of Membracidae are pre- comparative investigation of vibrational signals of Ci- sented. Taxonomic status and relationships of the taxa cadina, trying to cover all the families, subfamilies and studied are discussed basing on bioacoustic characters. tribes represented in the fauna of Russia and adjacent The concept of Membracidae deriving from a group territories as far as possible. within Cicadellidae, being not a sister-group of this Recently I published data on Iassides (Aphrodinae family, expressed earlier, is corroborated by bioacous- sensu Hamilton [1975], i.e. including Deltocephalinae tic data. + Iassinae + Penthimiinae) and Cicadellides (Cicadel- linae s.l. + Typhlocybinae) [Tishechkin, 2000c, 2001]. ÐÅÇÞÌÅ. Ïðèâåäåíû îñöèëëîãðàììû è îïèñà- The present paper is the third in the series and the last, íèÿ âèáðàöèîííûõ ñèãíàëîâ 50 âèäîâ öèêàäîê èç concerning Membracoidea. It includes descriptions of ïîäñåìåéñòâ Idiocerinae, Macropsinae, Agalliinae, signals of species from the group of related subfami- Adelungiinae, Megophthalminae è Ulopinae (Homo- lies usually named Ulopides (Idiocerinae, Macropsi- ptera, Cicadellidae) è 3 âèäîâ Membracidae. Íà îñ- nae, Agalliinae, Adelungiinae, Megophthalminae, Ul- íîâàíèè áèîàêóñòè÷åñêèõ ïðèçíàêîâ îáñóæäàåòñÿ opinae, etc.) and also, of several species of Mem- òàêñîíîìè÷åñêèé ñòàòóñ è ðîäñòâåííûå ñâÿçè èçó- bracidae. ÷åííûõ òàêñîíîâ. Âûñêàçàííîå ðàíåå ìíåíèå î The number of articles, providing oscillograms of òîì, ÷òî Membracidae âåäóò ñâîå íà÷àëî îò îäíîé acoustic signals of representatives of one or another of èç ãðóïï Cicadellidae, à íå ÿâëÿþòñÿ ñåñòðèíñêîé subfamilies studied is quite different. Several genera ãðóïïîé ýòîãî ñåìåéñòâà, ïîäòâåðæäàåòñÿ äàííû- of Macropsinae (Macropsis, Oncopsis, Hephathus) ìè áèîàêóñòèêè. are rather well studied due to recent taxonomic works in which signals analysis was involved [Claridge, Howse, Introduction 1968; Claridge, Reynolds, 1973; Claridge, Nixon, 1986; Tishechkin, 1992, 2000a, 2000b, 2002]. On the other The results of the first study of acoustic signals in hand, only data on two and three species respectively small Auchenorrhyncha (Cicadina with the exception are available on Agalliinae and Membracidae [Shaw, of singing cicadas — Cicadidae) were published by F. 1976; Hunt, 1993, 1994; Strübing, 1999). Idiocerinae, 12 D.Yu. Tishechkin

Megophthalminae and Ulopinae were never studied chirps, series). Succession of similar or different syllables, or after Ossiannilsson [1949]. For this reason, only verbal even any more or less prolonged signal with complex structure descriptions of their signals and recordings made by having determinable initial and end parts usually referred as means of radiograph, which is a pen recorder designed phrase. Succession of undeterminable duration, not changing in for registration of messages in Morse code, exist structure distinctly throughout its length and consisting of uniform pulses, repeating with constant period, is a trill. [Ossiannilsson, 1949]. All these works are discussed in The terms for signals elements listed above are used only in more details after descriptions of respective taxa. obvious cases, when the fragment may be classified with certain- Methods of recording and analysis of vibrational ty as pulse, syllable or trill. In dubious and intermediate situations signals are described in Tishechkin [2000c]. Only ex- I use neutral terms, such as “component”, “part”, etc. planations of main bioacoustic terms used in the paper are given herein. For a long time analysis of Cicadina signals was Calling signals. Signals produced by male ready to copu- used for discrimination between closely related spe- lation for attracting receptive female. Male can produce signals cies or for elucidation of the taxonomic status of of this type both being single and in the presence of other dubious forms only. Recently I undertook an attempt to individuals. Usually, male remains motionless and does not use acoustic characters for solving of taxonomic prob- demonstrate any activity after producing the signal, unless receptive female presents on the same stem. lems on subfamilies and tribes level in this group Reply signals. Signals produced by receptive female in [Tishechkin, 2000c, 2001]. The main problems existing reply to calling male. In most species, after certain time span in systematics of higher taxa of Ulopides concern the virgin female starts singing spontaneously. status of Adelungiinae (separate subfamily or the tribe of Courtship signals. Signals produced by male courting Agalliinae) and the position of Megophthalminae and female. The term is used only for signals differing distinctly in Ulopinae. Also, the supposition was expressed recently, temporal pattern from calling ones. that Membracidae, which were regarded as a sister- Copulatory signals. Signals emitted by male in last mo- group of Cicadellidae, indeed are specialized lineage of ments before copulation and/or at the moment of joining of leafhoppers related with Ulopinae, Megophthalminae genitalia. As in the previous case, the term is used only if the signal differs from calling and courtship ones. There issome confusion and Agalliinae [Rakitov, 1998; Dietrich et al., 2001]. in literature in using of this and previous terms. Occasionally, in Such conclusions concerning these problems, as can be species lacking courtship signals in a strict sense, signals made basing on bioacoustic data available, are given produced immediately before copulation are referred as court- after descriptions of signals in the respective groups. ship ones. In species, having both courtship and copulatory signals in their repertoire, the latters sometimes are named as Family Cicadellidae second type of courtship signals, courtship signals II, etc. Territorial signals. Usually produced by stationary sitting Subfamily Idiocerinae male either in the presence of individuals of the same or opposite sex, or spontaneously. As a rule, no apparent change in insect Idiocerus lituratus (Fallén, 1806) behaviour is observed either before or after the signals were Figs 1–3. produced. For this reason, these signals sometimes may be hardly distinguished from calling ones. However, territorial LOCALITY. Moscow Area, Voskresensk District, environs of signals for the most part have more simple and inconstant Beloozerskiy Town, from Salix cinerea L., 4.VIII.2000. Calling temporal pattern and sometimes are similar in different species. signals of 1 # are recorded at 23–24°C. Moreover, they never evoke response reaction of receptive SIGNALS. Calling signal is a long phrase having duration up female. Evidently, signals of this type serves for regulation of the to 20–25 s and consisting of two parts. The first part is a distance between individuals on the plant. succession of pulses, repeating with a period about 300–600 ms. Rivalry signals. As a rule, emitted by male in reply to calling Usually low-amplitude additional pulses are distinguishable on or rivalry signals of other individuals. Apparently, male usually oscillograms between high-amplitude main ones (Fig. 2). The produces signals of this type to drown calling of competing males. second part consists of syllables, repeating with period 1–1.3 s. In contrast with territorial ones, these signals were never regis- Each syllable includes several rather low-frequency pulses, tered from single males. Occasionally, several males sitting close similar with ones from the first part of signal, and ends with short to each other on the same stem start producing rivalry signals one having considerably higher carrier frequency (Fig. 3). For spontaneously. For this reason, sometimes there is no any well- this reason terminal pulses in syllables look distinctly darker on defined difference between this and previous types of signal. oscillograms at high speed. Present in repertoire of some Macropsinae and Adelungiinae only. In most part of other species studied male responds to calling Idiocerus stigmaticalis Lewis, 1834 of another one by producing calling or territorial signals. Figs 4–9. Call of distress. Under experimental conditions as a rule was registered from individuals got in some narrow place (in a LOCALITY. Moscow Area, environs of Bolshevo Town, corner of a cage, between a twig and a wall of a cage, etc.) and from Salix fragilis L., 16.VII.1984. Signals of 2 ## are recorded trying to escape from there, or touched by hand. at 23–24°C. Pulse is a brief elementary fragment of signal (or SIGNALS. Calling signal is a succession of syllables lasting up succession of sine waves) with fast increase and following to 15–20 s. In general, it is similar with the second part of calling of decrease of amplitude, i.e. separated from similar fragments the previous species. Each syllable consists of rather variable main by amplitude minimums. part and short abrupt click. The click has more high carrier Repeated with constant intervals uniform groups of similar frequency than the main part of syllable. Occasionally, low- or different in structure pulses are referred as syllables (= rolls, amplitude trill precedes or follows the signal (Figs 5–6). Vibrational communication in leafhoppers from Ulopides group 13

1

23 2 s 2

200 ms 3

200 ms 4

7 2 s 5

8 2 s 6

9 2 s 7

200 ms 8

200 ms 9

200 ms 10

12 2 s 11

13 2 s 12

200 ms 13

200 ms

Figs 1–13. Oscillograms of calling signals of Idiocerus lituratus (Fall.) (1–3), I. stigmaticalis Lew. (4–9) and Populicerus laminatus (Flor) (10–13). Faster oscillograms of the parts of signals indicated as “2–3”, “7–9” and “12–13” are given under the same numbers. Ðèñ. 1–13. Îñöèëëîãðàììû ïðèçûâíûõ ñèãíàëîâ Idiocerus lituratus (Fall.) (1–3), I. stigmaticalis Lew. (4–9) è Populicerus laminatus (Flor) (10–13). Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “2–3”, “7–9” è “12–13”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 14 D.Yu. Tishechkin

Populicerus laminatus (Flor, 1861) role in generation of signal. Apparently, low-frequen- Figs 10–13. cy vibrations presenting on oscillograms are resulted from these movements. LOCALITY. Moscow Area, Serpukhov District, environs of Tremulation (vibration by the parts of body, usual- Luzhki Village, from Populus tremula L., 21.VI.2000. Signals of 3 ly, by abdomen without contact with substrate) is one ## are recorded at the temperature 19–21°C. SIGNALS. Calling signal is a short phrase with duration of the most archaic and non-specialised mechanisms averaging 1.5–3 s. Its main component is a succession of pulses of acoustic communication in insects. For this rea- grouped in pairs, occasionally in the beginning of succession son, Idiocerinae seem to be the least advanced group pulses are indistinct (Figs 12–13). After it 2–4 short low- among the studied taxa of Ulopides. This fact corrob- amplitude pulses, separated by gaps about 150–250 ms follow orates the point of view of Emelyanov and co-authors (Fig. 12). Usually, low-frequency vibrations in the main part of [Anufriev, Emelyanov, 1988; Emelyanov, Kirillova, signal are clearly visible on oscillograms. Male vibrates by 1989], who place this subfamily in the beginning of abdomen distinctly when singing; low-frequency component of the list of Ulopides in their works. signal evidently results from abdomen movements. As a rule, male produces 2–5 phrases with intervals 0.5–1.5 s and then stops singing for several minutes. Two males sitting on the same Subfamily Macropsinae twig can sing simultaneously, so that their signals alternate. No other forms of competition behaviour were observed in this Pediopsis tiliae (Germar, 1831) species. Figs 26–45.

Populicerus populi (Linnaeus, 1761) LOCALITY. Moscow Area, Serpukhov District, environs of Luzhki Village, from small-leaved lime (Tilia cordata Miller), 14– Figs 14–21. 17.VI.1989. Signals of 11 ## are recorded at the temperature 22– 24°C. LOCALITIES. 1. Moscow, “Sokol’niki” Park. 17.VII.1992. Sig- SIGNALS. Calling signal is a single or repeating phrase nals of 2 ## are recorded at the temperature 25–26°C. consisting of three different parts (Figs 26–30). The first part 2. Moscow Area, environs of Bolshevo Town, from P. consists of rather variable pulses or syllables and lasts from 2– tremula, 11–12.VII.1984. Signals of 12 ## are recorded at 3 s up to several minutes (Figs 31–34). The second one is 21–22°C. composed of partially merged pulses repeating at a rate about SIGNALS. As in the previous species, calling signal is a 40/s, duration of this trill is about 1–1.5 s (Figs 32–34, 35–37). phrase, consisting of two successions of pulses. Usually, male The third part is a succession of discrete pulses following each produces several phrases repeating with a period about 2–5 s other with a period of 0.5–1 s. Number of pulses in it varies from (Figs 14–15). Occasionally, low-frequency trill precedes the 4–5 up to 20–30 (Figs 35–37). phrase described above (Figs 16–19). It can have rather small I have never registered female reply signals in P. tiliae. duration of 2–3 s, or, on the contrary, averages 10–20 s and Moreover, in this species, this is female, who walks in different more. Such more complex signal as a rule was registered from directions along the twig before mating, searching for male, male sitting in the same cage with female, but male never while the latter remains stationary, producing calling signals. demonstrated courtship behaviour during or after singing. Only after female approaches male within a distance of about Similarly with P. laminatus, low-frequency components of 3–4 cm, he suddenly runs to her and starts copulation. During signal are evidently emitted not only by timbals, but also by mounting female he produces copulatory signal, which is a tremulation movements. succession of pulses (Figs 38–41). Several males occurring on the same twig produce short Single male during grooming or not demonstrating any irregular trills of pulses during walking from time to time (Figs specific activity produces territorial signals, consisting of vari- 20–21). Maybe, these are territorial signals. able in structure irregular pulses (Figs 42–43). Several males sitting on the same twig occasionally can Populicerus confusus (Flor, 1861) react on each other’s calling by rivalry signals, which are Figs 22–25. irregular trills of short pulses (Figs 44–45). NOTE. Brief description of vibrational signals and mating LOCALITY. Moscow Area, environs of Pushkino, from Salix behaviour of this species is given in Tishechkin [1994]. cinerea L. on Ucha River bank. 14, 17.VII.1984. Signals of 6 ## are recorded at 23–25°C. Pediopsoides kurentsovi (Anufriev, 1977) SIGNALS. In general structure calling is similar with this in Figs 46–49. two other Populicerus species, but it has somewhat more complex temporal pattern due to additional succession of pulses in the middle of a signal. Also, more prolonged form of calling LOCALITY. The Southern Maritime Province, Pogranichny District, environs of Barabash-Levada Village, from Juglans with additional trill preceding the main part of signal, similar with mandshurica Maxim.; 1 #. 21.VII.1995. Shade air temperature — this in P. populi was heard, but not recorded in this species. As 27°C. in two previous species, a train of low-frequency pulses in the SIGNALS. Calling signal of this species is a long single beginning of signal evidently is emitted both by timbals and phrase lasting for about 20 s. It consists of several successions abdomen tremulation. of syllables, each syllable includes 4–6 low-amplitude pulses and one high-amplitude one. In the end of each succession All Idiocerinae studied distinctly vibrate by abdo- several additional discrete pulses present. men during singing. The mass of abdomen and ampli- NOTE. Interpretation of species is accepted after Hamilton tude of its movements is much higher, than in timbals, [1980]. In Anufriev, Emelyanov [1988] the species is mentioned so tremulation movements certainly play substantial under the name P. juglans Mats. Vibrational communication in leafhoppers from Ulopides group 15

14 2 s 15 15 200 ms

16

18 2 s 17

19 2 s 18

200 ms 19 200 ms 20 2 s 21 21 200 ms 22 2 s 23 24 23

200 ms 24

kHz 200 ms 7 6 5 4 3 2 1 0 25 200 ms

Figs 14–25. Vibrational signals of Populicerus populi (L.) (14–21), and P. confusus (Flor) (22–25): 14–15 — oscillograms of short form of calling signal of P. populi, 16–19 — same, long form, 20–21 — oscillograms of trills of pulses of males of P. populi, 22–24 — oscillograms of calling signal of P. confusus, 25 — same, oscillogram and sonogram. Faster oscillograms of the parts of signals indicated as “15”, “18–19”, “21” and “23–24” are given under the same numbers. Ðèñ. 14–25. Âèáðàöèîííûå ñèãíàëû Populicerus populi (L.) (14–21), è P. confusus (Flor) (22–25): 14–15 — îñöèëëîãðàììû êîðîòêîé ôîðìû ïðèçûâíîãî ñèãíàëà P. populi, 16–19 — òî æå, ïîëíàÿ ôîðìà, 20–21 — îñöèëëîãðàììû òðåëåé ïóëüñîâ ñàìöîâ P. populi, 22–24 — îñöèëëîãðàììû ïðèçûâíûõ ñèãíàëîâ P. confusus, 25 — òî æå, îñöèëëîãðàììà è ñîíîãðàììà. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “15”, “18–19”, “21” è “23–24”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 16 D.Yu. Tishechkin

26

28 6 s 27

29 6 s 28

32 35 2 s 29

2 s 33 36 30

31 2 s 34 37 31

200 ms 32

200 ms 33

200 ms 34

200 ms 35

200 ms 36

200 ms 37

200 ms

Figs 26–37. Oscillograms of calling signals of Pediopsis tiliae (Germ.). Faster oscillograms of the parts of signals indicated as “28– 29” and “31–37” are given under the same numbers. Ðèñ. 26–37. Îñöèëëîãðàììû ïðèçûâíûõ ñèãíàëîâ Pediopsis tiliae (Germ.). Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “28–29” è “31–37”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. Vibrational communication in leafhoppers from Ulopides group 17

38

39 40 41 2 s 39 200 ms 40

200 ms

41 200 ms

42

43 2 s 43

200 ms 44

45 2 s 45 200 ms

46 2 s 47 48 49 47

200 ms 48

200 ms 49

200 ms

Figs 38–49. Oscillograms of vibrational signals of Pediopsis tiliae (Germ.) (38–45) and Pediopsoides kurentsovi (Anufr.) (46–49): 38 — P. tiliae, calling and copulatory signals, 39 — same, calling signal, 40–41 — same, copulatory signal, 42–43 — same, territorial signals, 44–45 — same, rivalry signals, 46–49 — calling signal of P. kurentsovi. Faster oscillograms of the parts of signals indicated as “39–41”, “43”, “45” and “47–49” are given under the same numbers. Ðèñ. 38–49. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâ Pediopsis tiliae (Germ.) (38–45) è Pediopsoides kurentsovi (Anufr.) (46–49): 38 — P. tiliae, ïðèçûâíûé è êîïóëÿöèîííûé ñèãíàëû, 39 — òî æå, ïðèçûâíûé ñèãíàë, 40–41 — òî æå, êîïóëÿöèîííûé ñèãíàë, 42– 43 — òî æå, òåððèòîðèàëüíûé ñèãíàë, 44–45 — òî æå, ñèãíàë ñîïåðíè÷åñòâà, 46–49 — ïðèçûâíûé ñèãíàë P. kurentsovi. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “39–41”, “43”, “45” è “47–49”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 18 D.Yu. Tishechkin

Oncopsis flavicollis (Linnaeus, 1761) Oncopsis carpini (J. Sahlberg, 1871) Figs 50–61. Figs 72–76.

LOCALITIES. 1. Moscow Area, Serpukhov District, Pushchino- LOCALITY. Crimea, environs of Pereval’noe Village (halfway na-Oke Town, from common birch (Betula pendula Roth), 5– from Simferopol to Alushta), from Carpinus betulus L. 21.VI.1997. 8.VI.1985. Signals of more than 15 ## are recorded at 20–21°C. Signals of 3 ## are recorded at 24-26°C. 2. Altai Mountains, N shore of Teletskoe Lake, Altaiskiy SIGNALS. Calling signal in general is similar with these of Nature Reserve, environs of Yaylyu Village, from common birch. two previous species, but differs both in repetition period and 8.VII.1999. Calling signals of 1 # are recorded at the temperature shape of pulses (Figs 72–74). Rivalry signal is similar with these 30–35°C. of O. flavicollis and O. tsejensis (Figs 75–76). SIGNALS. Calling signal is a long single phrase, usually lasting from 10–15 up to 40–50 s (Figs 50–53). It begins with monotonous buzzing trill as a rule averaging several seconds. Oncopsis alni (Schrank, 1801) In the main part of signal trill is regularly (every 1–2 s) Figs 77–87. interrupted with short fragments of another kind (Figs 54–55). Their carrier frequency is always distinctly higher, than in the LOCALITY. Moscow Area, Serpukhov District, Pushchino-na- Oke Town, from black alder (Alnus glutinosa (L.) Gaertn.), 19–20, trill (Figs 56–57), but the amplitude varies greatly, depending 24.VI.1985. Signals of more than 10 individuals of both sexes are on physical characteristics of substrate (twig or stem on which recorded at 20–21°C. the insect sings). Signal ends with a short train of discrete SIGNALS. Calling signal is a long, monotonous, occasion- pulses. ally interrupting trill followed by succession of syllables, repeat- If male going along the twig contacts with another one sitting ing at a rate of about 2–3/s (Figs 77–80). When mounting on his way, one individual or both of them as a rule produce rivalry female, or sitting on her back during courtship, male produces signals (Figs 58, 60). Also, male emits signals of this type in reply copulatory signal, which is continuous trill (Figs 81–82). Terri- to calling or rivalry signal of another one, sitting on the same stem torial signals are rather variable syllables, following each other (Figs 51, 59). Rivalry signal is a long trill; sometimes it also with irregular intervals (Fig. 83). Rivalry signals are similar with includes buzzing fragments of varying length. these in other Oncopsis species (Figs 84–86). Also, both males Occasionally, male produces separate syllables with irreg- and females produce call of distress, when they are seized or ular intervals, not demonstrating any specific activity (Fig. 61). jammed, for instance in a narrow place between a twig and a Evidently, these are territorial signals. wall of a cage (Fig. 87). NOTES. Claridge and Nixon [1986], basing on both mor- phological and acoustic characters, came to the conclusion, that Oncopsis tristis (Zetterstedt, 1840) O. flavicollis sensu lato in Britain includes three closely related Figs 88–91. biological species. However, they did not give any scientific names to these forms, merely referring them as O. flavicollis LOCALITIES. 1. Moscow Area: (1) environs of Bolshevo Town, 1, 2 and 3. All the material studied by me, both from Moscow from common birch, 27.VI.1984, signals of 2 ## are recorded at Area and Altai Mts. belongs to O. flavicollis 2. the temperature 21°C; (2) environs of Pushkino Town, from common birch, 29–30.VI, 6.VII.1984, signals of 9 ## are recorded at the temperature 21–22°C Oncopsis subangulata (J. Sahlberg, 1871) 2. Altai Mountains, N shore of Teletskoe Lake, Altaiskiy Figs 62–66. Nature Reserve, environs of Yaylyu Village, from common birch together with O. flavicollis. 5.VII.1999. Calling signals of 1 # are LOCALITY. Moscow Area, environs of Pirogovo, the bank of recorded at the temperature 29°C. Klyaz’ma River, from birch (Betula sp.): 17.VI.1985, signals of 2 SIGNALS. Calling signal is a long phrase, lasting up to 40– ## are recorded at the temperature 21°C; 10.VI.1988, signals of 50 s. In general structure it is similar with signals of O. 2 ## are recorded at the temperature 25°C. subangulata, O. tsejensis and O. carpini, but differs in the SIGNALS. Calling is a long phrase, occasionally averaging shape of pulses and syllables. Temporal pattern of signals of more than a minute (Figs 62–64). It consists of alternating trills individuals from different populations is identical. of two types. As in O. flavicollis, relation of their amplitudes varies from signal to signal (Figs 65–66). Trills of both types are Acoustic signals of British Oncopsis species were composed of pulses or short syllables, but differ in the shape of studied by Claridge and co-authors (Claridge, Howse, pulses. Occasionally, pulses repetition rate somewhat decreas- 1968, Claridge, Reynolds, 1973, Claridge, Nixon, es before the end of signal. NOTE. Temporal pattern of calling signals in specimens 1986). In the first two papers only signals produced by studied is similar with this of songs of British ones, presented on male during mounting female before copulation were oscillograms in Claridge, Nixon [1986]. studied. Authors refer these signals as courtship ones, but according to terminology used in the present paper Oncopsis tsejensis Tishechkin, 1992 these are rather copulatory signals. Copulatory signal Figs 67–71. of O. tristis is a long trill similar with this of O. alni [Claridge, Howse, 1968]. In species from O. flavicollis LOCALITY. North Caucasus, North Ossetia, Tsey Canyon group (O. flavicollis, O. avellanae Edwards, 1920, O. (Ardon River basin) in the environs of Nizhniy Tsey Village subangulata, O. carpini) these signals are quite sim- (1700 m above sea level), from alder (Alnus incana (L.) Moench). ilar and consist of repeating phrases [Claridge, Rey- 27.VII.1990. Signals of 2 ## are recorded at the temperature 19– 20°C. nolds, 1973]. Nevertheless, basing on signals analysis, SIGNALS. As in the previous species, calling signal is a authors came to the conclusion that all four forms are long phrase, consisting of alternating fragments with different separate species. waveform (Figs 67–69). Rivalry signals are similar with these More detailed study of British Oncopsis species of O. flavicollis (Figs 70–71). from O. flavicollis group in Britain was provided by Vibrational communication in leafhoppers from Ulopides group 19

50 4 s 51

59 4 s 52 2 s 54 53

55 2 s 54 200 ms 55 200 ms kHz 7 6

5 4 3 2 1 0 56 200 ms 200 ms 57

60 2 s 58 200 ms 59

200 ms 60 200 ms

Figs 50–61. Vibrational signals of Oncopsis flavicollis (L): 50, 52–55 — oscillograms of calling signals, 51 — same, calling and rivalry signals, 56–57 — oscillograms and sonograms of calling signals, 58–60 — oscillograms of rivalry signals, 61 — oscillogram of territorial signal. Faster oscillograms of the parts of signals indicated as “54–55” and “59–60” are given under the same numbers. Ðèñ. 38–49. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâ Oncopsis flavicollis (L): 50, 52–55 — îñöèëëîãðàììû ïðèçûâíûõ ñèãíàëîâ, 51 — òî æå, ïðèçûâíûé ñèãíàë è ñèãíàë ñîïåðíè÷åñòâà, 56–57 — îñöèëëîãðàììû è ñîíîãðàììû ïðèçûâíûõ ñèãíàëîâ, 58–60 — îñöèëëîãðàììû ñèãíàëîâ ñîïåðíè÷åñòâà, 61 — îñöèëëîãðàììà òåððèòîðèàëüíîãî ñèãíàëà. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “54–55” è “59–60”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 20 D.Yu. Tishechkin

62 7 s 63 63

65 2 s 64 2 s 66 65 200 ms 66 200 ms 67

68 69 2 s 68 200 ms 69 200 ms 70

71 2 s 71 200 ms 72

73 74 2 s 73 200 ms 74

200 ms 75

2 s 76 76 200 ms

Figs 62–76. Oscillograms of vibrational signals of Oncopsis subangulata (J. Sahlb.) (62–66), O. tsejensis Tish. (67–71) and O. carpini (J. Sahlb.) (72–76): 62–66 — calling signals of O. subangulata, 67–69 — same, O. tsejensis, 70–71 — rivalry signals of O. tsejensis, 72–74 — O. carpini, calling signals, 75–76 — same, calling and rivalry signals. Faster oscillograms of the parts of signals indicated as “63”, “65–66”, “68–69”, “71”, “73–74” and “76” are given under the same numbers. Ðèñ. 62–76. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâ Oncopsis subangulata (J. Sahlb.) (62–66), O. tsejensis Tish. (67–71) è O. carpini (J. Sahlb.) (72–76): 62–66 — ïðèçûâíûå ñèãíàëû O. subangulata, 67–69 — òî æå, O. tsejensis, 70–71 — ñèãíàëû ñîïåðíè÷åñòâà O. tsejensis, 72–74 — O. carpini, ïðèçûâíûå ñèãíàëû, 75–76 — òî æå ïðèçûâíûå ñèãíàëû è ñèãíàëû ñîïåðíè÷åñòâà. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “63”, “65–66”, “68–69”, “71”, “73–74” è “76”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. Vibrational communication in leafhoppers from Ulopides group 21

77

2 s 79 78 2 s 80 79 200 ms 80 200 ms 81 2 s 82 82 200 ms 83 200 ms 84 2 s 85 86 85 200 ms 86 200 ms 87 200 ms 88 2 s 90 89 2 s 91 90 200 ms

91 200 ms

Figs 77–91. Oscillograms of vibrational signals of Oncopsis alni (Schrk.) (77–87) and O. tristis (Zett.) (88–91): 77–80 — O. alni, calling signals, 81–82 — same, copulatory signals, 83 — same, territorial signals, 84–86 — same, rivalry signals, 87 — same, call of distress, 88–91 — calling signals of O. tristis. Faster oscillograms of the parts of signals indicated as “79–80”, “82”, “85–86” and “90– 91” are given under the same numbers. Ðèñ. 77–91. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâ Oncopsis alni (Schrk.) (77–87) è O. tristis (Zett.) (88–91): 77–80 — O. alni, ïðèçûâíûå ñèãíàëû, 81–82 — òî æå, êîïóëÿöèîííûå ñèãíàëû, 83 — òî æå, òåððèòîðèàëüíûå ñèãíàëû, 84–86 — òî æå, ñèãíàëû ñîïåðíè÷åñòâà, 87 — òî æå, ñèãíàë ïðîòåñòà, 88–91 — ïðèçûâíûå ñèãíàëû O. tristis. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “79– 80”, “82”, “85–86” è “90–91”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 22 D.Yu. Tishechkin

Claridge and Nixon [1986]. Authors subdivided O. 2. Crimea, environs of Pereval’noe Village (halfway from flavicollis into three forms differing in structure of Simferopol to Alushta), from P. nigra L. 16.VI.1997. Signals of 2 both calling and copulatory signals and also, in some ## are recorded at 21–22°C. 3. Volgograd Area, Ilovlya River about 5 km from the mouth, morphological characters. Only one of these forms (so from P. nigra. 6–7.VI.1996. Signals of 5 ## are recorded at 18 called O. flavicollis 2) was found in Russia until now. and 21–22°C. Oscillograms of calling signals of O. flavicollis, O. 4. Orenburg Area, Sakmara River 20–25 km NNE of Kuvandyk subangulata and O. tsejensis are also presented in Town, environs of Churaevo Village, from P. nigra. 29.VI.1996. Tishechkin [1992]. Very brief descriptions of signals Signals of 2 ## are recorded at 27°C. SIGNALS. As in the previous species, there are two of 4 species are given in Tishechkin [1994]. forms of calling signals in M. graminea. Reduced form consists of three different parts (Figs 105–108), in full calling The genus Macropsis Lewis, 1834. two more parts similar with these in the signal of M. vicina Calling signals of 39 species are described in my present (Figs 109–116). Single male usually (but not always) produces reduced signals, whereas in the presence of female review of Macropsis from the territory of Russia he as a rule emits full ones. [Tishechkin, 2000a, 2002]. In the present paper only Territorial signal is a continuous trill consisting of pulses signals of selected species are described with the similar with these in the first part of calling, but following with purpose to illustrate as far as possible all the diversity longer and less regular intervals (Figs 117–118). of signals structure in the genus. Oscillograms show- NOTES. In two other closely related species dwelling on ing variability of signals temporal pattern in each spe- Populus (M. fuscinervis (Boh., 1845) and M. suspecta cies for the most part are not given. Geographical Tishechkin, 1994) calling signals structure is quite similar with variability of signals structure was not observed in the this in species described above. species of the genus. For this reason oscillograms of signals of the same species from different localities Macropsis cerea (Germar, 1837) also are not presented. Figs 119–126. LOCALITIES. 1. Moscow Area. Several geographical points in Macropsis vicina (Horvath, 1897) central and southern parts. Signals of more than 15 ## collected Figs 92–104. from different species of willows were recorded at the temperature 21–26°C. LOCALITIES. 1. Moscow Area, Serpukhov District, Pushchino- 2. East of Saratov Area, 10 km from Ozinki Town towards na-Oke Town, from Populus alba L. 3.VII.1985. Signals of 5 ## Ural’sk, from Salix vinogradovii Skvortsov, 1 #. 24.VI.1996. are recorded at 21°C. Recording was made at the shade air temperature 28–30°C. 2. Volgograd Area, Ilovlya River about 5 km from the mouth, 3. Orenburg Area, Sakmara River near Churaevo Village, 20– from P. alba. 10.VI.1996. Signals of 2 ## are recorded at 26–29°C. 25 km NNE of Kuvandyk, from Salix triandra L., 1 #. 29.VI.1996. 3. North-west Caucasus, Krasnodar Province, valley of Sukko Recording was made at the shade air temperature 26–27°C. River 12 km S of Anapa. 30.VI — 1.VII.1997. Calling signals of 4. Southern Tuva, environs of Erzin Village, from Salix 2 ## collected from P. alba are recorded at 27–28°C. ledebouriana Trautv. 30.VII.1989. Signals of 1 # were recorded at SIGNALS. There are two forms of calling signal in this the temperature 25°C. species. The first (reduced) one is a succession of syllables, 5. Amur Area, approximately 30 km W of Svobodny, from Salix lasting for approximately 10–20 s (Figs 92–94). Syllables at the udensis Trautv. et Mey., 1 #. 6.VII.1995. Recording was made at the beginning of signal have duration about 200–300 ms, then shade air temperature 25°C. several much longer ones (up to 1.5–2 s) follow. Their duration 6. The Southern Maritime Province, Pogranichny District, Komissarovka River near Barabash-Levada Village, from S. udensis, gradually decrease towards the end, so that initial and end 1 #. 13.VII.1995. Recording was made at the shade air temperature syllables in a signal are almost equal in length. 25°C. Calling of the second (full) type also begins with a succes- SIGNALS. Calling signal consists of repeating phrases, sion of syllables, but the last of the longer syllables suddenly following each other with a period of 2–4 s (Figs 119–122). Also, changes into monotonous buzzing fragment after which a train male emits the same signal on all stages of courtship behaviour of alternating short and long pulses follows (Figs 95–98). Single and immediately before copulation (Figs 123–124). male can produce signals of both types, but when courting Single male during walking and occasionally also when sitting female he produces only full calling. After emitting one or motionless produces continuous signal with highly varying pa- several such signals male mounts female and produces copula- rameters sounding like irregularly interrupting buzzing noise tory one which is an end part of calling, i.e. the buzzing fragment (Figs 125–126). Evidently, this is a territorial signal. and the train of pulses (Figs 99–102). In all cases observed copulation attempts were unsuccessful. Macropsis prasina (Boheman, 1852) Male, being single or in the presence of other individuals, Figs 127–133. also produces territorial signals, consisting of short syllables, as in the beginning of calling, but following with greater intervals LOCALITIES. 1. Moscow Area, environs of Pushkino (about (Figs 103–104). Occasionally, he can sing in such a manner for 15 km from the north-eastern boundary of Moscow), from Salix half an hour and more. cinerea L. on the bank of Ucha River. 23, 25.VI.1984 and 24.VI.1988. Signals of 8 ## are recorded at the temperature 20– Macropsis graminea (Fabricius, 1798) 21 and 27°C. 2. Northern part of Saratov Area, environs of Khvalynsk, near Figs 105–118. Ulyanino Village, from S. cinerea. 15, 17.VI.1996. Signals of 5 ## are recorded at the temperature 23 and 33°C. LOCALITIES. 1. Moscow Area, Serpukhov District, Pushchino- 3. East of Saratov Area, 10 km E of Ozinki Town towards na-Oke Town, from Populus nigra var. pyramidalis Spach. 27– Ural’sk, from S. cinerea. 24.VI.1996. Signals of 3 ## are recorded 28.VI.1985. Signals of 3 ## are recorded at 21–22°C. at the temperature 27–30°C. Vibrational communication in leafhoppers from Ulopides group 23

92 2 s 93 94 93 200 ms 94 200 ms

95

96 4 s 96

2 s 97 98 97 200 ms 98 200 ms 99

6 s 100 101 100 2 s 101

102 2 s 102 200 ms

103

104 200 ms 104

200 ms

Figs 92–104. Oscillograms of vibrational signals of Macropsis vicina (Horv.): 92–94 — reduced form of calling signal, 95–98 — same, full form, 99 — calling and copulatory signals, 100 — calling signal, 101–102 — copulatory signals, 103–104 — territorial signals. Faster oscillograms of the parts of signals indicated as “93–94”, “96–98”, “100–102” and “104” are given under the same numbers. Ðèñ. 92–104. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâ Macropsis vicina (Horv.): 92–94 — ïðèçûâíûå ñèãíàëû, ðåäóöèðîâàííàÿ ôîðìà, 95–98 — òî æå, ïîëíàÿ ôîðìà, 99 — ïðèçûâíûé è êîïóëÿöèîííûé ñèãíàëû, 100 — ïðèçûâíûé ñèãíàë, 101–102 — êîïóëÿöèîííûå ñèãíàëû, 103–104 — òåððèòîðèàëüíûå ñèãíàëû. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “93–94”, “96–98”, “100– 102” è “104”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 24 D.Yu. Tishechkin

105

2 s 106

107 108 2 s 107

200 ms 108 200 ms 109 110 111 4 s 110

2 s 111 2 s

112 2 s 113 114 115 116 113 200 ms 114 200 ms 115

200 ms 116

200 ms

117

118 2 s 118

200 ms

Figs 105–118. Oscillograms of vibrational signals of Macropsis graminea (F.): 105–108 — calling signals, reduced form, 109–116 — same, full form, 117–118 — territorial signals. Faster oscillograms of the parts of signals indicated as “107–108”, “110–111”, “113– 116” and “118” are given under the same numbers. Ðèñ. 105–118. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâ Macropsis graminea (F.): 105–108 — ïðèçûâíûå ñèãíàëû, ðåäóöèðîâàííàÿ ôîðìà, 109–116 — òî æå, ïîëíàÿ ôîðìà, 117–118 — òåððèòîðèàëüíûå ñèãíàëû. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “107– 108”, “110–111”, “113–116” è “118”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. Vibrational communication in leafhoppers from Ulopides group 25

119

121 2 s 120 2 s 122 121 200 ms 122 200 ms

123

124 2 s 124 200 ms 125

126 2 s 126 200 ms 127

128 2 s 128 200 ms 129 2 s 130 200 ms 131 200 ms 132

133 2 s 133 200 ms

Figs 119–133. Oscillograms of vibrational signals of Macropsis cerea (Germ.) (119–126) and M. prasina (Boh.) (127–133): 119–122 —M. cerea, calling signals, 123–124 — same, signals, produced by male immediately before copulation, 125–126 — same, territorial signals, 127–128 —M. prasina, calling signals, 129–131 — same, copulatory signals, 132–133 — same, calling and rivalry signals. Faster oscillograms of the parts of signals indicated as “121–122”, “124”, “126”, “128” and “133” are given under the same numbers. Ðèñ. 199–133. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâ Macropsis cerea (Germ.) (119–126) è M. prasina (Boh.) (127–133): 119– 122 — M. cerea, ïðèçûâíûå ñèãíàëû, 123–124 — òî æå, ñèãíàëû, èçäàâàåìûå ñàìöîì íåïîñðåäñòâåííî ïåðåä êîïóëÿöèåé, 125– 126 — òî æå, òåððèòîðèàëüíûå ñèãíàëû, 127–128 — M. prasina, ïðèçûâíûå ñèãíàëû, 129–131 — òî æå, êîïóëÿöèîííûå ñèãíàëû, 132–133 — òî æå, ïðèçûâíûé ñèãíàë è ñèãíàë àãðåññèè. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “121–122”, “124”, “126”, “128” è “133”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 26 D.Yu. Tishechkin

4. Orenburg Area, Sakmara River near Churaevo Village, 20– SIGNALS. Calling signal is a succession of syllables, 25 km NNE of Kuvandyk Town, from Salix viminalis L., 2 ##. repeating with a period 300–600 ms (Figs 146–149). Total 29.VI.1996. Shade air temperature — 27–28°C. length of signal averages 5–20 s. 5. Altai Mountains, S shore of Teletskoe Lake, Chiri, mouth Several males sitting on the same twig can answer to one of Kyga River, from S. viminalis. 11.VII.1999. Calling signals of 1 another’s calling by short successions of pulses, which are # are recorded at the temperature 22°C. SIGNALS. Calling signal consists of short phrases usually rivalry signals (Figs 150–151). Territorial signal is similar with repeating with a period about 3–8 s (Figs 127–128). Each phrase these in other willow-dwelling species and is a long trill with begins with monotonous trill, sounding like buzz, in the end part irregular temporal pattern (Figs 152–153). of it 2–4 fragments with another waveform present. Then several discrete syllables follow. Other males sitting on the Macropsis haupti Wagner, 1941 same twig as a rule produce rivalry signals in reply to calling one Figs 154–159. (Figs 132–133). These signals are successions of syllables, same as in the second part of calling. LOCALITIES. 1. NW Caucasus, Krasnodar Area, Sukko River Male produces the same signal both being single and sitting in the environs of Sukko Village 12 km South of Anapa, from Salix next to female during courtship. After mounting female he emits elbursensis Boiss. 30.VI.1997. Calling signals of 1 # were recorded another signal with quite different temporal structure (Figs 129– at 26°C. 131). Undoubtedly, this is copulatory one, but in several cases 2. Volgograd Area, Ilovlya River 5–7 km from the mouth, from Salix vinogradovii Skvortsov. 10.VI.1996. Calling signals of 2 ## it was also registered from single individual. were recorded at 22–25 and 29–30°C. SIGNALS. Calling signal is a phrase consisting of rather Macropsis verbae Anufriev, Zhiltsova, 1982 low-amplitude trill, after which a succession of high-amplitude Figs 134–139. syllables follows (Figs 154–157). Duration of both parts and of phrase in total varies greatly. Territorial signal is similar with LOCALITIES. 1. Moscow Area, Serpukhov District, Pushchino- these in other species (Figs 158–159). na-Oke Town, from Salix acutifoliaWilld. on the bank of Oka River. 2.VII.1985. Signals of 4 ## are recorded at 21°C. Macropsis ocellata Provancher, 1872 2. Volgograd Area, Ilovlya River about 5–7 km from the mouth, from S. acutifolia. 10.VI.1996. Signals of 3 ## are recorded at 31– Figs 160–167. 32°C. SIGNALS. Calling signal consists of two different parts LOCALITIES. 1. Moscow Area, Serpukhov District, environs of (Figs 134–137). The first part is a monotonous or modulated Luzhki Village, Salix alba L. on the bank of Oka River. 16.VII.1985 fragment lasting up to 5–8 s (Fig. 136), the second one consists and 30.VII.1994. Signals of 6 ## were recorded at 22–23°C. of short repeating phrases (Fig. 137). Sometimes singing male 2. North-west Caucasus, 12 km S of Anapa, Sukko River, from S. alba, 4 ##. 2.VII.1997. Shade air temperature — 29–31°C. omits the first part of signal. 3. Astrakhan’ Area, Dosang Railway Station, from S . alba Territorial signal is irregular buzzing trill (Figs 138–139). (possibly, S. alba x S. fragilis) in a flood-land between Volga and Akhtuba rivers, 7.VII.2000. Signals of 2 ## were recorded at 26°C. Macropsis gravesteini Wagner, 1953 4. Orenburg Area, Sakmara River near Churaevo Village, 20– Figs 140–145. 25 km NNE of Kuvandyk Town, from S. alba, 5 ##. 29.VII.1996. Shade air temperature — 24–28°C. LOCALITIES. 1. Crimea, environs of Pereval’noe Village SIGNALS. Full calling signal consists of train of short monot- halfway from Simferopol to Alushta, from Salix alba L. 16– onous fragments, sounding like continuous regularly interrupting 17.VI.1997. Calling signals of 3 ## are recorded at the buzz, after which a trill of partially merged pulses follows (Figs 160, temperature 22–25°C. 162, 164). Total length of signal in some cases exceeds 30–40 s. 2. Saratov Area, environs of Khvalynsk Town, near Ulyanino Sometimes male omits the second part of signal (trill of pulses) (Figs Village, from S. alba and S. fragilis L. 15, 17.VI.1996. Signals of 5 161, 165). Occasionally, inverse situation takes place and the first ## are recorded at 31 and 22°C. part may be almost entirely reduced, but the second one remains 3. Volgograd Area, Ilovlya River about 5–7 km from the mouth, well-pronounced (Figs 163, 166–167). from S. alba. 8.VI.1996. Calling signals of 3 ## are recorded at During the only observed attempt of copulation male emit- 20–26°C. ted calling several times, sitting close to female. Then he SIGNALS. Calling signal is a succession of short phrases, mounted her back does not producing any signals at that usually repeating with a period about 2–5 s (Figs 140–143). moment. In several seconds female rejected male. Territorial signal is a prolonged highly variable trill (Figs 144– 145). Male emits signals of this type both being single and in Macropsis fuscula (Zetterstedt, 1828) reply to calling of another individual. Figs 168–173.

Macropsis leporina Tishechkin, 1997 LOCALITIES. 1. Moscow Area: (a) environs of Serpukhov Figs 146–153. Town; (b) environs of Bronnitsy Town. Signals of more than 10 specimens collected fromRubus idaeus L., R. caesius L. and R . nessensis LOCALITIES. 1. Chita Area, Ingoda River 15 km E of Urul’ga W. Hall were recorded at the temperature 24–26°C. (Karymskiy District), from Salix schwerinii E.Wolf, 7 ##. 29.VI — 2. Southern Kazakhstan, Almaty and Zailiyskiy Alatau Mt. 2.VII.1995. Shade air temperature — 25–27°C. Ridge in the environs of the city, from R. caesius. 3.VII.1994. Signals 2. Amur Area, 30 km W of Svobodny, from Salix udensis Trautv. of 4 ## were recorded at the temperature 30–31°C. et Mey. 7–8.VII.1995. Signals of 4 ## are recorded at the SIGNALS. Calling signal consists of short phrases repeating temperature 21–22 and 27°C. with a period from 1–2 up to 4–5 s (Figs 168–170). Two males 3. The Southern Maritime Province, Pogranichny District, sitting on the same stem sometimes sing simultaneously, so that environs of Barabash-Levada, from S. udensis. 13.VII.1995. Signals phrases produced by different individuals alternate (Fig. 171). I of 1 # are recorded at the temperature 25°C. have never registered rivalry signals in this species. Vibrational communication in leafhoppers from Ulopides group 27

134

2 s

135

136 137 2 s 136

200 ms 137 200 ms 138

139 2 s 139

200 ms

140

142 2 s 141

143 2 s 142

200 ms

143 200 ms

144

145 2 s 145 200 ms

Figs 134–145. Oscillograms of vibrational signals of Macropsis verbae Anufr., Zhilts. (134–139) and M. gravesteini Wagn. (140– 145): 134–137 — M. verbae, calling signals, 138–139 — same, territorial signals, 140–143 — M. gravesteini, calling signals, 144–145 — same, territorial signals. Faster oscillograms of the parts of signals indicated as “136–137”, “139”, “142–143” and “145” are given under the same numbers. Ðèñ. 134–145. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâ Macropsis verbae Anufr., Zhilts. (134–139) è M. gravesteini Wagn. (140– 145): 134–137 — M. verbae, ïðèçûâíûå ñèãíàëû, 138–139 — òî æå, òåððèòîðèàëüíûå ñèãíàëû, 140–143 — M. gravesteini, ïðèçûâíûå ñèãíàëû, 144–145 — òî æå, òåððèòîðèàëüíûå ñèãíàëû. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “136–137”, “139”, “142–143” and “145”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 28 D.Yu. Tishechkin

146 2 s 148 147 2 s 149 148 200 ms 149 200 ms 150 2 s 151 151 200 ms

152 2 s 153 153 200 ms 154 2 s 156 155

2 s 157 156 200 ms 157 200 ms

158

159 2 s 159 200 ms

Figs 146–159. Oscillograms of vibrational signals of Macropsis leporina Tish. (146–153) and M. haupti Wagn. (154–159): 146– 149 — M. leporina, calling signals, 150–151 — same, rivalry signals, 152–153 — same, territorial signals, 154–157 — M. haupti, calling signals, 158–159 — same, territorial signals. Faster oscillograms of the parts of signals indicated as “148–149”, “151”, “153”, “156– 157” and “159” are given under the same numbers. Ðèñ. 146–159. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâ Macropsis leporina Tish. (146–153) è M. haupti Wagn. (154–159): 146– 149 — M. leporina, ïðèçûâíûå ñèãíàëû, 150–151 — òî æå, ñèãíàëû ñîïåðíè÷åñòâà, 152–153 — òî æå, òåððèòîðèàëüíûå ñèãíàëû, 154–157 — M. haupti, ïðèçûâíûå ñèãíàëû, 158–159 — òî æå, òåððèòîðèàëüíûå ñèãíàëû. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “148–149”, “151”, “153”, “156–157” è “159”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. Vibrational communication in leafhoppers from Ulopides group 29

160

162 4 s 161

165 4 s 162 2 s 164 163

166 167 2 s 164 200 ms 165 200 ms 166

200 ms 167

200 ms 168

169 170 2 s 169

200 ms 170

200 ms 171

2 s 172

173 2 s 173 200 ms

Figs 160–173. Oscillograms of vibrational signals of Macropsis ocellata Prov. (160–167) and M. fuscula (Zett.) (168–173): 160– 167 — M. ocellata, different types of calling signals, 168–170 — M. fuscula, calling signal, 171 — same, calling alternation of two males, 172–173 — same, territorial signals. Faster oscillograms of the parts of signals indicated as “162”, “164–167”, “169–170” and “173” are given under the same numbers. Ðèñ. 160–173. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâ Macropsis ocellata Prov. (160–167) è M. fuscula (Zett.) (168–173): 160– 167 —M. ocellata, ðàçíûå òèïû ïðèçûâíîãî ñèãíàëà, 168–170 — M. fuscula, ïðèçûâíûé ñèãíàë, 171 — òî æå, ÷åðåäóþùèåñÿ ïðèçûâíûå ñèãíàëû äâóõ ñàìöîâ, 172–173 — òî æå, òåððèòîðèàëüíûå ñèãíàëû. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “162”, “164–167”, “169–170” è “173”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 30 D.Yu. Tishechkin

Territorial signal is a prolonged succession of pulses with 6. Kazakhstan, Almaty and foothills of Zailiyskiy Alatau highly variable temporal structure (Figs 172–173). Mountain Ridge in the environs of the city, from Elaeagnus sp. 2.VII.1994. Signals of 5 ## are recorded at 31°C. Macropsis scutellata (Boheman, 1845) SIGNALS. Calling signal is a succession of syllables Figs 174–178. following each other with a period of about 2–4 s (Figs 193– 194). Copulatory signal is also a succession of syllables, but LOCALITIES. 1. Moscow Area, Voskresensk District, environs their temporal pattern is quite different (Figs 195–197). The of Beloozerskiy Town, from Urtica dioica L. 29.VII.1991. Signals latter signal male normally produces when sitting on female’s of one # were recorded at the temperature 24°C. back immediately before attempt of copulation, but occasion- 2. North-west Caucasus, Krasnodar Province, environs of ally it was also registered from single individual. Call of Praskoveevka Village 15 km S of Gelendzhik, from U. dioica, 1 #. distress is a train of pulses (Figs 197–198). Sitting stationary, 12.VII.1997. Shade air temperature — 26°C. especially during grooming, male produces short trains of SIGNALS. Calling signal consists of two parts with differ- pulses from time to time (Figs 199–200). Evidently, these are ent temporal pattern (Figs 174–176). Its duration usually aver- territorial signal. ages 20–30 s. Territorial signal is similar with this of the previous species (Figs 177–178). Macropsis pictipes (Horvath, 1904) Macropsis megerlei (Fieber, 1868) Figs 201–206. Figs 179–187. LOCALITIES. 1. Southern Tuva, environs of Erzin Village, LOCALITY. Moscow Area, Serpukhov District, environs of flood-land of Erzin River, fromHippophae rhamnoidesL. 21.VII.1989. Luzhki Village, 12 and 16.VII.1990. Signals of 8 ## collected from Signals of 2 ## are recorded at 24°C. Rosa majalis Herrm. were recorded at 22–23°C. 2. Kazakhstan, environs of Almaty, Zailiyskiy Alatau Moun- SIGNALS. Calling signal consists of alternating fragments tain Ridge, from H. rhamnoides near the river. 30.VI.1994. Signals of two different types, one of which is about twice shorter, than of 1 # are recorded at 31°C. another (Figs 179–182). As a rule, after one long fragment 1– SIGNALS. Calling signal is a long phrase averaging up to 3 short ones follow. These signals were registered both from 40–50 s and consisting of trill followed by succession of syllables single male and from one sitting close to female. It was female, (Figs 201–206). Sometimes male omits the trill and produces the who was searching for the mate, whereas singing male was train of syllables only. Two males singing on the same twig sitting stationary in my experiments. When female approached emitted syllables alternately, but never produced rivalry signals male within a distance about 1–2 cm, he suddenly ran towards in my experiments. her and mounted her back. After that he emitted copulatory signal and started copulation. Copulatory signal begins with a trill, then a succession of alternating syllables of two types Macropsis glandacea (Fieber, 1868) follows (Figs 183–185). Figs 207–214. Territorial signals in this species are continuous trains of syllables or single pulses with highly variable structure (Figs LOCALITIES. 1. North-west Caucasus, 12 km S of Anapa, Sukko 186–187). River, from Ulmus carpinifolia Gled. 1.VII.1997. Signals of 1 # are recorded at 27–29°C. Macropsis idae Emeljanov, 1964 2. North-east Caucasus, Chechnya, Terskiy Mountain Ridge Figs 188–192. in the environs of Grozny Town, from U. carpinifolia. 27.VI.1986. Signals of 4 ## are recorded at the temperature 24°C. LOCALITY. Signals of 6 ## collected from Rosa sp. in the SIGNALS. Calling signal is a long succession of syllables Botanical Garden in Almaty (Kazakhstan) were recorded at (Figs 207–210). The same signal male emits sitting on female’s 3.VII.1994 at the temperature 32°C. back immediately before copulation (Figs 211–212). Territorial SIGNALS. Calling signal consists of syllables repeating signal is a long trill (Figs 213–214). with a period of 1–1.5 s (Figs 188–191). Also, long trills of discrete pulses were registered in males of this species (Figs Macropsidius abrotani Emeljanov, 1964 188–189, 192). Male produces this signal either spontaneously or in reply to calling of another one. Thus, these trills act both as Figs 215–228. territorial and rivalry signal. LOCALITIES. 1. Moscow Area, Serpukhov District, from Arte- Macropsis elaeagni Emeljanov, 1964 misia abrotanum L. on Oka River bank in the environs of Luzhki Figs 193–200. Village, 11–12.VII.1985. Signals of more than 10 specimens of both sexes are recorded at the temperature 21–22°C. LOCALITIES. 1. Moscow, from ornamental trees of Elaeagnus 2. East of Saratov Area, 10–15 km E of Ozinki Town. A. angustifolia L. in the park near the Moscow State University. 14, abrotanum on roadside in steppe. 22.VI.1996. Signals of 1 # are 24.VII.1998. Signals of 7 ## are recorded at 25 and 29°C. recorded at the temperature 28–29°C. 2. Crimea, Kerchenskiy Peninsula, E shore of Kazantipskiy SIGNALS. Calling signal is a long single phrase, lasting for Bay, Zolotoe Village, from cultivated E. angustifolia. 27.VI.1997. 20–30 s and more and consisting of three different successions Signals of 5 ## are recorded at the temperature 30–31°C. of syllables (Figs 215–223). The shape of syllables depends 3. North-west Caucasus, 12 km S of Anapa, Sukko River, from much on physical characteristics of substrate on which the E. angustifolia. 3.VII.1997. Signals of 5 ## are recorded at 27–28°C. insect sings (Figs 220–222 and oscillograms on Fig. 223). As 4. North Caucasus, Chechnya, Terskiy Mountain Ridge in the in other Macropsinae, male also produces this signal in close environs of Grozny Town, from E. angustifolia. 19–22.VI.1986. proximity with female. Only after mounting female he starts Signals of 10 ## are recorded at the temperature 22–24°C. 5. Southern Urals, Guberlya River near Guberlya railway emitting copulatory signal, differing in structure from calling station, 25 km W of Orsk, from E. angustifolia. 5.VII.1996. Signals one (Figs 224–225). All observed copulation attempts were of 5 ## are recorded at 25–30°C. unsuccessful. Vibrational communication in leafhoppers from Ulopides group 31

174

175 176 2 s 175 200 ms 176 200 ms 177

178 2 s 178 200 ms

179

181 2 s 180

182 2 s 181

200 ms 182 200 ms 183

184 185 2 s 184

200 ms 185 200 ms 186

187 2 s 187 200 ms

Figs 174–187. Oscillograms of vibrational signals of Macropsis scutellata (Boh.) (174–178) and M. megerlei (Fieb.) (179–187): 174– 176 — M. scutellata, calling signals, 177–178 — same, territorial signals, 179–182 — M. megerlei, calling signals, 183–185 — same, copulatory signals, 186–187 — same, territorial signals. Faster oscillograms of the parts of signals indicated as “175–176”, “178”, “181– 182”, “184–185” and “187” are given under the same numbers. Ðèñ. 174–187. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâ Macropsis scutellata (Boh.) (174–178) è M. megerlei (Fieb.) (179–187): 174–176 — ïðèçûâíûå ñèãíàëû M. scutellata, 177–178 — òî æå, òåððèòîðèàëüíûå ñèãíàëû, 179–182 — ïðèçûâíûå ñèãíàëû M. megerlei, 183–185 — òî æå, êîïóëÿöèîííûå ñèãíàëû, 186–187 — òî æå, òåððèòîðèàëüíûå ñèãíàëû. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “175–176”, “178”, “181–182”, “184–185” è “187”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 32 D.Yu. Tishechkin

188

190 2 s 189

2 s 191 192 190

200 ms 191 200 ms 192 200 ms 193

2 s 191 194 200 ms 195 2 s 196 196 200 ms

197

198 2 s 198 200 ms 199

200 2 s 200 200 ms

Figs 188–200. Oscillograms of vibrational signals of Macropsis idae Em. (188–192) and M. elaeagni Em. (193–200): 188–189 — M. idae, calling and rivalry signals, 190–191 — same, calling signals, 192 — same, rivalry signal, 193–194 — M. elaeagni, calling signals, 195–196 — same, copulatory signals, 197 — same, copulatory signals of one male and call of distress of another one; the former male attempts to copulate with the latter one, 198 — same, call of distress, 199–200 — same, territorial signals. Faster oscillograms of the parts of signals indicated as “190–192”, “194”, “196”, “198” and “200” are given under the same numbers. Ðèñ. 188–200. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâ Macropsis idae Em. (188–192) è M. elaeagni Em. (193–200): 188–189 — M. idae, ïðèçûâíûå ñèãíàëû è ñèãíàëû ñîïåðíè÷åñòâà, 190–191 — òî æå, ïðèçûâíûå ñèãíàëû, 192 — òî æå, ñèãíàëû ñîïåðíè÷åñòâà, 193–194 — M. elaeagni, ïðèçûâíûå ñèãíàëû, 195–196 — òî æå, êîïóëÿöèîííûå ñèãíàëû, 197 — òî æå, êîïóëÿöèîííûå ñèãíàëû îäíîãî ñàìöà è ñèãíàëû ïðîòåñòà äðóãîãî; ïåðâûé ñàìåö ïûòàåòñÿ êîïóëèðîâàòü ñî âòîðûì, 198 — òî æå, ñèãíàëû ïðîòåñòà, 199– 200 — òî æå, òåððèòîðèàëüíûå ñèãíàëû. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “190–192”, “194”, “196”, “198” è “200”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. Vibrational communication in leafhoppers from Ulopides group 33

201

202 203 4 s 202

204 205 2 s 203 2 s 206 204 200 ms 205 200 ms 206 200 ms 207

209 2 s 208

210 2 s 209 200 ms 210 200 ms 211

212 2 s 212

200 ms 213

214 2 s 214 200 ms

Figs 201–214. Oscillograms of vibrational signals of Macropsis pictipes (Horv.) (201–206) and M. glandacea (Fieb.) (207–214): 201–206 — calling signal of M. pictipes, 207–210 — M. glandacea, calling signals, 211–212 — same, signals, produced by male immediately before copulation, 213–214 — same, territorial signals. Faster oscillograms of the parts of signals indicated as “202–206”, “209–210”, “212” and “214” are given under the same numbers. Ðèñ. 201–214. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâ Macropsis pictipes (Horv.) (201–206) è M. glandacea (Fieb.) (207–214): 201–206 — ïðèçûâíûé ñèãíàë M. pictipes, 207–210 — M. glandacea, ïðèçûâíûå ñèãíàëû, 211–212 — òî æå, ñèãíàëû, èçäàâàåìûå ñàìöîì íåïîñðåäñòâåííî ïåðåä êîïóëÿöèåé, 213–214 — òî æå, òåððèòîðèàëüíûå ñèãíàëû. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “202–206”, “209–210”, “212” and “214”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 34 D.Yu. Tishechkin

215 4 s 216 216 2 s 218 219 217

220 221 222 2 s 218 200 ms 219 200 ms

220 kHz 200 ms221 200 ms 222 200 ms 7

6 5 4 3 2 1 0 223 200 ms 224

225 2 s 225 200 ms 226

227 2 s 227 200 ms 228 200 ms

Figs 215–228. Vibrational signals of Macropsidius abrotani Em.: 215–222 — oscillograms of calling signals, 223 — oscillograms and sonograms of three parts of calling signal, 224–225 — copulatory signal, 226–227 — calling and rivalry signals, 228 — call of distress. Faster oscillograms of the parts of signals indicated as “216”, “218–222”, “225” and “227” are given under the same numbers. Ðèñ. 215–228. Âèáðàöèîííûå ñèãíàëû Macropsidius abrotani Em.: 215–222 — îñöèëëîãðàììû ïðèçûâíûõ ñèãíàëîâ, 223 — îñöèëëîãðàììû è ñîíîãðàììû òðåõ ÷àñòåé ïðèçûâíîãî ñèãíàëà, 224–225 — êîïóëÿöèîííûé ñèãíàë, 226–227 — ïðèçûâíûé ñèãíàë è ñèãíàë ñîïåðíè÷åñòâà, 228 — ñèãíàë ïðîòåñòà. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “216”, “218–222”, “225” è “227”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. Vibrational communication in leafhoppers from Ulopides group 35

Usually, male produces rivalry signals in reply to calling of The genus Hephathus Ribaut, 1952. another individual singing nearby (Figs 226–227). Same signals were registered from several males sitting close to each other in Revision of species of this genus from the territory the same cage. Also, in female of this species call of distress of Russia was published recently [Tishechkin, 2000b]. similar with this in other Macropsinae was recorded (Fig. 228). Calling signals of males from two different localities do not The paper includes descriptions and oscillograms of differ in temporal structure. calling signals, on which signals of individuals from different localities are presented and variability of Macropsidius sahlbergi (Flor, 1861) their temporal pattern is shown. For this reason, only Figs 229–235. several most representative oscillograms for each Hephathus species are given in the present paper. LOCALITIES. Moscow Area: (1) environs of Zvenigorod Town, sandy slopes on bank of Moskva River near Lutsyno Village, from Hephathus nanus (Herrich–Schäffer, 1835) Artemisia campestris L. 27–28.VI.1988, signals of 4 ## are recorded at the temperature 26–28°C, 2.VII.1991, signals of 2 ## Figs 251–255. and 2 $$ are recorded at the temperature 30°C; (2) Serpukhov District, from A. campestris on Oka River bank in the environs LOCALITIES. 1. Crimea, mountains 3–4 km E of Pereval’noe of Luzhki Village, 30.VII.1993. Signals of 1 # are recorded at the Village (halfway from Simferopol to Alushta), dry meadow with temperature 25°C. steppe vegetation in the mountains. 18.VI.1997. Calling signals SIGNALS. Calling signal is a short phrase averaging in of 2 ## were recorded at the temperature 25–26°C. duration approximately 1–2 s. Single male can produce phrases 2. East of Saratov Area, about 15 km NE of Ozinki Town, dry with irregular intervals; after hearing female reply he starts ruderal vegetation on the roadside. 22.VI.1996. Calling signals of singing more actively and regularly, emitting signals with pauses 1 # were recorded at the temperature 33°C. about 0.5 s (Figs 229–231). 3. Southern Urals, Guberlinskiye Mountains in the environs Female reply is a short train of pulses (Figs 234–235). of Guberlya railway station 25 km W of Orsk, rather wet hollow Usually it somewhat overlaps with the end part of male calling with high herbaceous steppe vegetation between the hills. 8– 9.VII.1996. Calling signals of 2 ## were recorded at the or follows immediately after it. As in most other leafhoppers, temperature 26–28°C. male runs along the plant searching for female, whereas the SIGNALS. Calling signal is a successions of syllables, latter remains stationary during this alternate calling. After following each other almost without gaps, or separated by finding female male mounts her back, produces copulatory intervals up to 1–1.5 s. In the former case, signal usually lasts signal, lasting up to 20–30 s (Figs 232–233) and starts copula- up to 10 s (Figs 251, 253–254). If male produces syllables with tion. If copulation attempt appears to be unsuccessful, male rather long intervals, he can sing unceasingly for one minute and emits copulatory signal once more before the next one. more (Figs 252, 255). NOTES. In European Russia the species for a long time Macropsidius serratus Logvinenko, 1965 was confused with H. achilleae (see Tishechkin, 2000b). Figs 236–241. Signals of H. achilleae are presented under the name H. nanus in Tishechkin [1994]. LOCALITY. Crimea, Kerchenskiy Peninsula, E shore of Kazantipskiy Bay, environs of Zolotoe Village, from Artemisia sp. along salted dry river-bed. 25–26.VI.1997. Signals of 5 ## were Hephathus freyi (Fieber, 1868) recorded at 27–29°C. Figs 256–258. SIGNALS. Calling signal is a phrase consisting of initial fragment with irregular structure, sounding like buzz, and sev- LOCALITIES. 1. Rostov Area, Oblivskiy district, environs of eral syllables with higher amplitude (Figs 236–239). Normally, Sosnovy (=Oporny) Village on Chir River. 8.VIII.1991, male emits several phrases one after another; in the first one 17.VIII.1992. Calling signals of 4 ## were recorded at the initial part (buzz) can last for 20–30 s and more, in the successive temperature 27–30°C. phrases it is much shorter and does not exceed 2–3 s. Several 2. Volgograd Area, Ilovlya River about 5–7 km from the mouth. males sitting on the same plant can sing alternately, but some- 10.VI.1996. Calling signals of 1# were recorded at the temperature times one male produces rivalry signals in reply to calling of 25°C. 3. East of Saratov Area, about 15 km NE of Ozinki Town, dry another individual (Figs 240–241). ruderal vegetation on the roadside in the same biotope, where H. nanus was collected. 22.VI.1996. Calling signals of 1 # were Macropsidius niger (Matsumura, 1915) recorded at the temperature 31–32°C. Figs 242–250. 4. South-eastern Azerbaidzhan, Talysh Mountains, environs of Lerik Village. 10.VII.1987. Calling signals of 1 # were recorded at the temperature 20°C. LOCALITIES. 1. South Siberia, southern Tuva, environs of 5. Southern Turkmenistan, environs of Dushak Town. 9.V.1994. Erzin Village, from Artemisia glauca Pall. ex Willd. on the roadside. Calling signals of 1 # were recorded at the temperature 31°C. 11.VIII.1989. Signals of 3 ## were recorded at 27°C. 2. The Southern Maritime Province, Pogranichny District, SIGNALS. Calling signal is a trill of discrete pulses, environs of Barabash-Levada Village, from Artemisia sp. (Sect. repeating with a period about 90–160 ms. Additional low- Artemisia), 2 ##. 13.VII.1995. Shade air temperature — 26°C. amplitude pulses present between main ones in the beginning SIGNALS. Calling signal is a succession of short syllables of a signal. Overall duration of trill varies greatly and averages or pulses, repeating with a period 100–600 ms (Figs 242–248). from 1–2 up to 30–40 s. Duration of signal varies greatly averaging from 1.5–2 up to NOTES. From the territory of the former Soviet Union the 20–25 s in different cases. Also, in males call of distress was species was recorded under the name H. unicolor (Lindberg, registered (Figs 249–250). Calling signals of males from 1926), which is a junior synonym of H. freyi [Tishechkin, different localities are similar. 2000b]. 36 D.Yu. Tishechkin

229

230 2 s 231 230 200 ms 231

200 ms 232

233 2 s 233 $ $ 200 ms 234

$ 200 ms 235

200 ms

236

2 s 237

238 2 s 239 238

200 ms 239

200 ms 240

2 s 241 241

200 ms

Figs 229–241. Oscillograms of vibrational signals of Macropsidius sahlbergi (Flor) (229–235) and M. serratus Logv. (236–241): 229– 231 — M. sahlbergi, calling signals, 232–233 — same, copulatory signal, 234–235 — same, male calling and female reply, 236–239 — M. serratus, calling signals, 240–241 — same, calling and rivalry signals. Faster oscillograms of the parts of signals indicated as “230– 231”, “233”, “238–239” and “241” are given under the same numbers. Ðèñ. 229–241. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâ Macropsidius sahlbergi (Flor) (229–235) è M. serratus Logv. (236–241): 229–231 — M. sahlbergi, ïðèçûâíûå ñèãíàëû, 232–233 — òî æå, êîïóëÿöèîííûé ñèãíàë, 234–235 — òî æå, ïðèçûâ ñàìöà è îòâåò ñàìêè, 236–239 — M. serratus, ïðèçûâíûå ñèãíàëû, 240–241 — òî æå, ïðèçûâíûå ñèãíàëû è ñèãíàëû ñîïåðíè÷åñòâà. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “230–231”, “233”, “238–239” è “241”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 200 ms

Vibrational communication in leafhoppers from Ulopides group 37

242

2 s 243

246 2 s 244 2 s 247 245

2 s 248 246

200 ms 247 200 ms 248

200 ms 249

250 2 s 250 200 ms 251

253 254 2 s 252 2 s 255 253

200 ms 254

200 ms 255

200 ms

Figs 242–255. Oscillograms of vibrational signals of Macropsidius niger (Mats.) (242–250) and Hephathus nanus (H.-S.). (251– 255): 242–248 — M. niger, calling signals, 249–250 — same, call of distress, 251–255 — calling signals of H. nanus. Faster oscillograms of the parts of signals indicated as “246–248”, “250” and “253–255” are given under the same numbers. Ðèñ. 242–255. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâ Macropsidius niger (Mats.) (242–250) è Hephathus nanus (H.-S.). (251– 255): 242–248 — M. niger, ïðèçûâíûå ñèãíàëû, 249–250 — òî æå, ñèãíàëû ïðîòåñòà, 251–255 — ïðèçûâíûå ñèãíàëû H. nanus. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “246–248”, “250” è “253–255”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 38 D.Yu. Tishechkin

Hephathus achilleae Mityaev, 1967 Auchenorrhyncha: on the first stage male produces Figs 259–265. calling signals spontaneously, then receptive female starts singing in reply to his calling. For some time LOCALITIES. 1. Moscow Area, environs of Zvenigorod, dry sandy mates sing alternately, female being motionless, male slopes on the bank of Moskva River near Lutsyno Village. 27– running in different directions along the plant search- 28.VI.1988. Calling signals of 7 ## were recorded at 26–28°C. 2. North Caucasus, Chechnya, steppe on the slopes of Terskiy ing for her. After finding female male mounts her back, Mountain Ridge in the environs of Grozny. 19.VI.1986. Calling produces copulatory signal and starts copulation. signals of 2 ## were recorded at the temperature 23–24°C. In no Macropsinae courtship signals were regis- 3. East of Saratov Area, 10 km E of Ozinki Town towards tered. In all cases, when mating behaviour was ob- Ural’sk. 25.VI.1996. Calling signals of 2 ## were recorded at the served, male produced calling signals both being single temperature 30–31°C. 4. Southern Urals, Guberlinskiye Mountains in the environs and in close contact with female. Only when sitting on of Guberlya railway station 25 km W of Orsk, rather wet hollow the back of female in last moments before copulation, with high herbaceous steppe vegetation between the hills, together male usually produces signals of another type, namely, with H. nanus. 8.VII.1996. Calling signals of 2 ## were recorded copulatory ones. However, in some Macropsis species at the temperature 26–27°C. SIGNALS. Calling signal consists of syllables, having dura- male goes on producing calling even at that moment. In tion about 160–400 ms and repeating with irregular intervals several species of the genus copulatory signals occa- (Figs 259–263). Sometimes several discrete pulses are visible sionally were registered from single males. Maybe, on oscillograms in the initial part of syllable. Occasionally, this these were individuals on the stage of most high sexual part lengthens greatly, so that the whole duration of syllable activity. exceeds 1–2 s. Arboreal Macropsinae (genera Pediopsis, Pediop- Also, call of distress was registered in H. achilleae. As in soides, Oncopsis and Macropsis) as a rule have terri- most other species, it is a continuous train of pulses with torial signals in their acoustic repertoire. Single males irregular structure (Figs 264–265). usually produce signals of this type during walking or NOTES. See the same item in the description of signals of H. nanus. grooming (when scraping the body surface by legs and creating on it the coating of brochosomes — for details Some data on vibrational communication in Mac- see Rakitov, 1998) and before producing calling. More ropsinae were published earlier (Tishechkin, 1994), rarely signals of this type were registered from individ- very brief descriptions and oscillograms of signals of uals sitting motionless and demonstrating no specific several species are presented in this work. activity. In grass-dwelling species (Macropsidius, Hep- Acoustic communication system in various genera hathus) territorial signals were not recorded. of this subfamily is quite different. In representatives In all Macropsinae with the exception of Hep- of Pediopsis, Pediopsoides, Oncopsis and, also, in hathus rivalry signals were registered in appropriate poplar-feeding Macropsis species calling signals are situations. Male produces signals of this type in reply long single phrases sometimes having very complex to calling of another one. Also, several males sitting temporal pattern. In willow-dwelling Macropsis call- in the same cage can produce rivalry signals alternate- ing signals are either single or regularly repeating ly or simultaneously from time to time. Among Ci- phrases, the latter case is more abundant. In species of cadellidae such signals were also registered in Ade- the genus living on Rosaceae, Elaeagnaceae and plants lungiinae, in most other groups males demonstrating from other families except Salicaceae calling usually competition behaviour produce calling signals in turn. consists of short repeating phrases or syllables, still Also, in representatives of all genera studied calls their temporal pattern as a rule is rather complex. In the of distress were recorded both in males and in females. genus Macropsidius both long, complex phrases (M. Insects produce these signals in various stress situa- abrotani) and short, simple signals (M. sahlbergi, M. tions (being hold or got in some narrow place and trying niger) present. Calling signals of Hephathus are the to escape), or if male attempts to copulate with another simplest among Macropsinae. one or with female not ready to mating. Neither in P. tiliae, nor in several Macropsis It is possible, that some peculiar features of commu- species, in which mating behaviour was studied, fe- nication system in arboreal Macropsinae, such as absence male reply signals were registered. Moreover, in P. of female reply signals and well-developed territorial and tiliae and M. megerlei it was female, who walked rivalry behaviour are resulted from their rather sedentary along the plant searching for the mate, whereas sing- way of life. Males of Pediopsis, Macropsis and Oncop- ing male remained on the same place (in other cases sis can sit on the same place for many hours producing searching behaviour in female was not observed, be- calling with irregular intervals. In such situation territori- cause insects were sitting near each other before any al and rivalry signals may play important role in regula- of them started singing). Evidently, in these species tion of distance between individuals, because these in- female does not produce reply signals, and communi- sects leave long-occupied place only very reluctantly in cation between representatives of different sexes contrast with most of other leafhoppers, for instance, becomes one-way, as in singing cicadas (Cicadidae). Aphrodinae (= Deltocephalinae), Typhlocybinae, etc. In Macropsidius (M. sahlbergi) communication In actively moving insects female reply draws atten- system and mating behaviour are typical for small tion of male and thus prevents him from leaving the Vibrational communication in leafhoppers from Ulopides group 39

256 2 s

257 2 s 258 258 200 ms

259 2 s 261 260 2 s 263 262 261 200 ms 262 200 ms 263 200 ms 264 2 s 265 265 200 ms 266

268 2 s 267

269 2 s 268 200 ms 269 200 ms

Figs 256–269. Oscillograms of vibrational signals of Hephathus freyi (Fieb.) (256–258), H. achilleae Mit. (259–265) and Agallia brachyptera (Boh.) (266–269): 256–258 — calling signals of H. freyi, 259–263 — same of H. achilleae, 264–265 — call of distress of H. achilleae, 266–269 — calling signals of A. brachyptera. Faster oscillograms of the parts of signals indicated as “258”, “261–263”, “265” and “268–269” are given under the same numbers. Ðèñ. 256–269. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâ Hephathus freyi (Fieb.) (256–258), H. achilleae Mit. (259–265) è Agallia brachyptera (Boh.) (266–269): 256–258 — ïðèçûâíûå ñèãíàëû H. freyi, 259–263 — òî æå, H. achilleae, 264–265 — ñèãíàë ïðîòåñòà H. achilleae, 266–269 — ïðèçûâíûå ñèãíàëû A. brachyptera. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “258”, “261–263”, “265” è “268–269”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 40 D.Yu. Tishechkin plant on which receptive female occurs. In sedentary Anaceratagallia ribauti (Ossiannilsson, 1938) leafhoppers singing male does not change his location Figs 287–289. for a long time, so female can reach him being guided by his signals, not producing reply ones. LOCALITIES. Moscow Area, Serpukhov District, dry meadow In general, judging by such acoustic characters as on bank of Oka River near Luzhki Village, 31.VII.1993, signals of 1 # are recorded at the temperature 23–24°C. rather complex calling signals consisting of single or SIGNALS. Calling signal is similar with this of the previous repeated phrases (or complicated syllables) and well- species, but temporal structure of syllables in the second part of developed territorial and rivalry signals, Macropsinae phrase is quite different. seem to be most closely related with Agalliinae and Adelungiinae (see below). Anaceratagallia aciculata (Horvath, 1894) Figs 290–296. Subfamily Agalliinae LOCALITY. Transvolga part of Astrakhan’ Area, environs of Dosang Railway Station, flood–land between Akhtuba and Volga Agallia brachyptera (Boheman, 1847) rivers, on ruderal vegetation in irrigation ditch. 4.VII.2000. Signals Figs 266–282. of 2 ## are recorded at the temperature 26–27°C. SIGNALS. Calling signal is a very long phrase, occasionally LOCALITIES. Moscow Area: (1) environs of Pirogovo, wet having duration up to 1–2 minutes and more. As in two previous meadow on the bank of Klyaz’ma River, 30.VII.1987, signals of 2 species, it consists of two parts differing both in temporal pattern ## are recorded at the temperature 22°C; (2) environs of and syllables repetition period. Pushkino, meadow on the bank of Ucha River, 23.VII.1988, signals of 1 # are recorded at the temperature 27–28°C; (3) Voskresensk Anaceratagallia chalchica Dlabola, 1967 District, environs of Beloozerskiy Town, from Rumex acetosella L. in pine forest, 26.VII.1988, signals of 1 # are recorded at the Figs 297–299. temperature 26–27°C, 7.VII.1991, signals of 1 # and 2 $$ are recorded at the temperature 26–27°C. LOCALITY. South Siberia, southern Tuva, environs of Erzin SIGNALS. Calling signal is a short phrase lasting for 1–3 s Village, from Artemisia glauca Pall. ex Willd. on the roadside. (Figs 266–269). It consists of trill followed by short fragment in 11.VIII.1989. Signals of 1 # were recorded at 27°C. which pulses are indistinct (Figs 268–269). Frequency spectra SIGNALS. Calling signal is a phrase lasting for 5–6 s and of these two parts differ conspicuously, which is clearly audible consisting of more or less uniform pulses grouped in syllables. even by human ear (Fig. 274). Male can produce phrases with Temporal pattern of signal gradually changes towards the end, irregular intervals from 1–2 s up to several minutes (Fig. 266), in the end part of phrase boundaries between syllables become or sings ceaselessly for about half a minute or more (Fig. 267). indistinct. In the latter case pauses between phrases do not exceed 100– NOTES. Determination of species is based on Dlabola 200 ms. [1967] (original description) and Vilbaste [1980]. Female reply signal is a continuous trill of pulses, repeating with approximately the same rate as in the first part of male Austroagallia sinuata (Mulsant et Rey, 1855) calling (Figs 270–271). In my experiments female answered Figs 300–308. both to full calling and to its first part only (the trill of pulses). After hearing female reply male starts running along the plant LOCALITIES. (1) North Caucasus, Chechnya, Terskiy Moun- in different directions searching for the mate. His signals become tain Ridge in the environs of Grozny Town. 4.VII.1986. Signals more regular and long due to extending of the first part (Figs 272– of 1 # were recorded at 28°C. 273). Also, he flaps the wings several times at the end of each (2) Ukraine, Kherson Area, Sivash Lake, steppe on seashore, from phrase (Figs 273–274); thus calling signal transforms into Artemisia sp. 16.VIII.1986. Signals of 1 # were recorded at 28°C. SIGNALS. As in some other Cicadellidae species, there courtship one. After finding female male mounts her back. For are two forms of calling signal in A. sinuata. Usually, calling is several seconds he flaps wings rhythmically at a rate about 6– a short phrase with a duration about 1.5 s, consisting of two 8/s and produces copulatory signal, which is a trill (Figs 275– syllables, the first being at least three times as long as the 280), then partners join their genitalia. second. As a rule, male produces several phrases with intervals Male sitting motionless, both single and in the presence of of 1.5–4 s (Figs 300–301, 303–306). Occasionally, the last other individuals also produces short pulses following with irreg- phrase in the succession has rather long additional part, averag- ular intervals and sounding like clicks or ticking (Figs 281–282). ing 15–20 s and more (Figs 302, 307). Evidently these are territorial signals. Also, call of distress consisting of short irregular successions of pulses once was registered in male of this species (Figs 308). Anaceratagallia venosa (Fourcroy, 1785) Figs 283–286. Detailed study of acoustic signals and associated behaviour of two Agalliinae species, namely, Agallia LOCALITIES. Moscow Area, Voskresensk District, environs of constricta van Duzee and Agalliopsis novella (Say) was Beloozerskiy Town, from Rumex acetosella L. in pine forest, provided by Shaw [1976]. Both signals temporal struc- 26.VII.1991, signals of 2 ## are recorded at the temperature 26– ture and set of their functional types, as well as mating 27°C. SIGNALS. Calling signal is a long phrase lasting up to 20– behaviour in these species does not differ principally 30 s. It consists of two successions of syllables, differing both from these in ones involved in my study. Signals were in temporal pattern and syllables repetition period. Usually registered by microphone, for this reason the author phrases follow with irregular intervals from 1–2 seconds up to speaks of sounds, not of vibrational signals. Calling several minutes. signals of both species (referred as “common sounds”) Vibrational communication in leafhoppers from Ulopides group 41

270 2 s 271 $ 271

200 ms 272

2 s 273 273 kHz wing flaps 200 ms 7 6 5 4 3 2 1 0 274 200 ms 275

276 277 6 s 278 276 2 s 277 2 s 2 s 279 280 278 200 ms 280 200 ms 281

282 2 s 282

200 ms

Figs 270–282. Vibrational signals of Agallia brachyptera (Boh.): 270–271 — oscillograms of male calling and female reply signals, 272–273 — oscillograms of courtship signal, 274 — same, oscillogram and sonogram, 275 — oscillogram of courtship and copulatory signals, 276 — oscillogram of courtship signal, 277–280 — oscillograms of copulatory signal (277, 279 — wing flaps, 278, 280 — signal, produced by timbals), 281–282 — oscillograms of territorial signals. Faster oscillograms of the parts of signals indicated as “271”, “273”, “276–280” and “282” are given under the same numbers. Ðèñ. 270–282. Âèáðàöèîííûå ñèãíàëû Agallia brachyptera (Boh.): 270–271 — îñöèëëîãðàììû ïðèçûâíîãî ñèãíàëà ñàìöà è îòâåòíîãî ñèãíàëà ñàìêè, 272–273 — îñöèëëîãðàììû ñèãíàëà óõàæèâàíèÿ, 274 — òî æå, îñöèëëîãðàììà è ñîíîãðàììà, 275 — îñöèëëîãðàììà ñèãíàëà óõàæèâàíèÿ è êîïóëÿöèîííîãî ñèãíàëà, 276 — îñöèëëîãðàììà ñèãíàëà óõàæèâàíèÿ, 277–280 — îñöèëëîãðàììû êîïóëÿöèîííîãî ñèãíàëà (277, 279 — óäàðû êðûëüåâ, 278, 280 — ñèãíàë, èçäàâàåìûé ïðè ïîìîùè òèìáàëîâ), 281– 282 — îñöèëëîãðàììû òåððèòîðèàëüíûõ ñèãíàëîâ. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “271”, “273”, “276–280” è “282”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 42 D.Yu. Tishechkin

283

285 286 2 s 284 2 s 285 200 ms 286

200 ms 287

288 289 2 s 288 200 ms

289

200 ms 290

4 s 291

292 293 4 s 292

294 2 s 295 293 2 s 296 294

200 ms 295

200 ms 296

200 ms

Figs 283–296. Oscillograms of calling signals of Anaceratagallia venosa (Fourcroy.) (283–286), A. ribauti (Oss.) (287–289) and A. aciculata (Horv.) (290–296). Faster oscillograms of the parts of signals indicated as “285–286”, “288–289” and “292–296” are given under the same numbers. Ðèñ. 283–296. Îñöèëëîãðàììû ïðèçûâíûõ ñèãíàëîâ Anaceratagallia venosa (Fourcroy.) (283–286), A. ribauti (Oss.) (287–289) è A. aciculata (Horv.) (290–296). Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “285–286”, “288–289” è “292–296”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. Vibrational communication in leafhoppers from Ulopides group 43

297

298 299 2 s 298

200 ms 299

200 ms 300

303 304 2 s 301

305 2 s 302

306 307 2 s 303

200 ms 304

200 ms 305 200 ms 306

200 ms 307

200 ms 308

200 ms

Figs 297–308. Oscillograms of vibrational signals of Anaceratagallia chalchica Dlab. (297–299) and Austroagallia sinuata (M. R.). (300–308): 297–299 — calling signals of A. chalchica, 300–301 and 303–305 — calling signals of A. sinuata, short form, 302, 306– 307 — same, signal with additional part, 308 — same species, call of distress. Faster oscillograms of the parts of signals indicated as “298–299” and “303–307” are given under the same numbers. Ðèñ. 297–308. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâ Anaceratagallia chalchica Dlab. (297–299) è Austroagallia sinuata (M. R.). (300–308): 297–299 — ïðèçûâíûå ñèãíàëû A. chalchica, 300–301 è 303–305 — ïðèçûâíûå ñèãíàëû A. sinuata, êîðîòêàÿ ôîðìà, 302, 306–307 — òî æå, ñèãíàë ñ äîïîëíèòåëüíîé ÷àñòüþ, 308 — òîò æå âèä, ñèãíàë ïðîòåñòà. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “298–299” è “303–307”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 44 D.Yu. Tishechkin are long and complex phrases lasting for 15–20 s and signal varies from 4–5 up to 15–20 s. Other males sitting nearby more, and consisting of several parts with different on the same plant usually answer to calling by rivalry signals, temporal pattern. In A. constricta, the first part of calling consisting of syllables, similar in temporal pattern with these in sometimes lengthens greatly reaching duration up to 3– calling one, but following with less regular intervals (Fig. 318). 4 minutes and more; such transformed signal was re- Quite often several males sitting close to each other on the same ferred as “common sound II”. It must be noted, that twig start producing rivalry signals spontaneously (Figs 321– oscillograms of “common sound I” of A. constricta and 322). NOTES. The specimens recorded were identified by Prof. of “common sound” of A. novella on figs 1 and 3 A.F. Emelyanov (St.-Petersburg). respectively in the paper cited are overturned, so that the zero must be shifted to the right end of the time scale. In both species receptive female answers male Symphypyga repetekia Kusnezov, 1929 calling producing reply signals (“female common Figs 323–327. sounds”, according to terminology used in the paper). LOCALITY. Transvolga part of Astrakhan’ Area, environs of For some time insects sing alternately, during this duet Dosang Railway Station, from Calligonum aphyllum (Pall.) Gürke male calling becomes shorter, its temporal structure in sand desert. 6.VII.2000. Signals of 5 ## are recorded at the somewhat changes, thus calling transforms into court- temperature 26–27°C. ship signal. Female reply also changes slightly, so the SIGNALS. Calling signal is a succession of short syllables, author refers this modified signal as “female courtship somewhat similar with this of A. nigronervosus (Figs 323– sound”. In last seconds before copulation male produc- 325). Shape of syllables sometimes slightly changes from the es rather variable trill of pulses — copulatory signal (= beginning to the end of signal. As in most Adelungiinae species, “precopulatory sound”). In A. novella male also vi- other males answer to calling signal by rivalry ones (Figs 326– brates by wings at this moment. 327). Also, both sexes of both species produce short pulses or clicks repeating with irregular intervals, so Platyproctus tessellatus Lindberg, 1924 called “ticking sounds”. Signals of this type occur in Figs 328–333. many different behavioural contexts, both in motion- less, spaced individuals and preceding or following any LOCALITY. Transvolga part of Astrakhan’ Area, environs of Dosang Railway Station, from Calligonum aphyllum (Pall.) Gürke kind of activity, including walking, jumping, producing in sand desert. 30.VI, 5.VII.2000. Signals of more than 10 ## are of calling signal, etc. Signals of the same kind I have recorded at the temperature 25–26°C. registered in A. brachyptera (see above). SIGNALS. Spontaneously produced calling signal includes 2–4 phrases, following each other with a period 0.5–3 s (Figs Subfamily Adelungiinae 329, 332–333). Each phrase consists of low-amplitude succes- sion of partly merged pulses and 1–3 high-amplitude ones, sounding like abrupt clicks. Several males usually sang alterna- Achrus albicosta (Kusnezov, 1929) tively (Fig. 329), but never produced territorial or rivalry signals Figs 309–316. in my experiments. Male courting female sings more regularly and continuously LOCALITY. Southern Turkmenistan, the road Dushak — (Figs 328, 330–331), running along the plant in different directions Tedzhen 30 km East of Dushak: (1) from Haloxylon aphyllum searching for the mate. Phrases of the signal at these moments (Minkw.) Iljin, 27.V.1990, signals of 3 ## are recorded at the become much longer, than in spontaneous calling due to elonga- temperature 31°C; (2) from Calligonum sp., 10.V.1994, signals of 1 # are recorded at the temperature 31°C. tion of the first (low-amplitude) part. Then male mounts female SIGNALS. Calling signal is a succession of short phrases, and starts copulation. I have neither registered, nor even heard following each other with a period about 1.5–3 s. Shape of pulses female reply signals. Still, there is no doubt, that they present in on oscillograms sometimes differs much in different signals acoustic repertoire of the species, because in all cases, when depending on physical characteristics of substrate and relative copulation was observed, it was male, who started searching for position of singing insect and vibrotransducer (Figs 309, 311 and female, while the latter sat motionless. Evidently, reply signals 310, 312). As in some Macropsinae species, male can answer the were not discernible through the noises produced by moving calling of another one by rivalry signals, consisting of short trains male due to their low amplitude. of discrete pulses (Figs 313–314). Territorial signals produced by male either being single or in the presence of other individuals Melicharella callifrons Mitjaev, 1971 consist of trills with irregular structure (Figs 315–316). Figs 334–337. NOTES. The specimens recorded were identified by Prof. A.F. Emelyanov (St.-Petersburg). LOCALITY. Transvolga part of Astrakhan’ Area, environs of Dosang Railway Station, from Calligonum aphyllum (Pall.) Gürke Achrus nigronervosus Lindberg, 1924 in sand desert. 15.VII.2000. Signals of 5 ## are recorded at the Figs 317–322. temperature 28°C. SIGNALS. Calling signal is a short syllable (Figs 334–336). LOCALITY. Southern Turkmenistan, the road Dushak — In my experiments males produced signals with irregular inter- Tedzhen 30 km East of Dushak, from Calligonum sp., 27.V.1990. vals. Other males presenting on the same stem as a rule answer Signals of 3 ## are recorded at the temperature 27°C. to calling signal emitting continuous irregular rivalry ones (Figs SIGNALS. Calling signal is a succession of short syllables 334, 337). following each other at a rate 5–6/s. Succession ends with a NOTES. Determination of species is based on Mityaev fragment having irregular structure (Figs 317–320). Duration of [1971]. 200 ms

Vibrational communication in leafhoppers from Ulopides group 45

309

311 2 s 310

312 2 s 311

200 ms 312 200 ms 313

314 2 s 314 200 ms 315

316 2 s 316

200 ms 317

2 s 319 318

320 2 s 319

200 ms 320

200 ms 321 2 s 322 322 200 ms

Figs 309–322. Oscillograms of vibrational signals of Achrus albicosta (Kusn.) (309–316) and A. nigronervosus Lindb. (317–322): 309–312 — A. albicosta, calling signals, 313–314 — same, calling and rivalry signals, 315–316 — same, territorial signals, 317, 319– 320 — A. nigronervosus, calling signals, 318 — same, calling and rivalry signals, 321–322 — same, rivalry signals. Faster oscillograms of the parts of signals indicated as “311–312”, “314”, “316”, “319–320” and “322” are given under the same numbers. Ðèñ. 309–322. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâ Achrus albicosta (Kusn.) (309–316) è A. nigronervosus Lindb. (317–322): 309–312 — A. albicosta, ïðèçûâíûå ñèãíàëû, 313–314 — òî æå, ïðèçûâíûå ñèãíàëû è ñèãíàëû ñîïåðíè÷åñòâà, 315–316 — òî æå, òåððèòîðèàëüíûå ñèãíàëû, 317, 319–320 — A. nigronervosus, ïðèçûâíûå ñèãíàëû, 318 — òî æå, ïðèçûâíûå ñèãíàëû è ñèãíàëû ñîïåðíè÷åñòâà, 321–322 — òî æå, ñèãíàëû ñîïåðíè÷åñòâà. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “311–312”, “314”, “316”, “319– 320” è “322”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 46 D.Yu. Tishechkin

323

324 325 2 s 324 200 ms 325 200 ms 326

327 2 s 327 200 ms 328

2 s 330 331 329 2 s 332 333 330 200 ms 331 200 ms 332

200 ms 333 200 ms 334

336 337 2 s 335 336 200 ms 200 ms 337 200 ms

Figs 323–337. Oscillograms of vibrational signals of Symphypyga repetekia Kusn. (323–327), Platyproctus tessellatus Lindb. (328– 333) and Melicharella callifrons Mit. (334–337): 323–325 — S. repetekia, calling signals, 326–327 — same, rivalry signals, 328– 333 — calling signals of P. tessellatus, 334 — M. callifrons, calling and rivalry signals, 335–336 — same, calling signals, 337 — same, rivalry signals. Faster oscillograms of the parts of signals indicated as “324–325”, “327”, “330–333” and “336–337” are given under the same numbers. Ðèñ. 323–337. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâ Symphypyga repetekia Kusn. (323–327), Platyproctus tessellatus Lindb. (328– 333) è Melicharella callifrons Mit. (334–337): 323–325 — S. repetekia, ïðèçûâíûå ñèãíàëû, 326–327 — òî æå, ñèãíàëû ñîïåðíè÷åñòâà, 328–333 — ïðèçûâíûå ñèãíàëû P. tessellatus, 334 — M. callifrons, ïðèçûâíûé ñèãíàë è ñèãíàëû ñîïåðíè÷åñòâà, 335–336 — òî æå, ïðèçûâíûå ñèãíàëû, 337 — òî æå, ñèãíàëû ñîïåðíè÷åñòâà. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “324–325”, “327”, “330–333” è “336–337”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. Vibrational communication in leafhoppers from Ulopides group 47

Adelungiinae form a well-defined group very closely Subfamily Ulopinae related to Agalliinae and are regarded either as a tribe of this subfamily (for instance, Emelyanov, 1975) or as Utecha trivia (Germar, 1821) a separate group [Davis, 1975; Al-Ne’amy, Linnavuori, Figs 346–358. 1982; Rakitov, 1998, etc.]. Davis [1975] divides it into two tribes, Achrini and Adelungiini; among the genera LOCALITY. North Caucasus, Chechnya, Terskiy Mountain studied the first two ones (Achrus and Symphypyga) Ridge in the environs of Grozny Town. 9.VII.1986. Signals of 7 ## are recorded at the temperature 30–32°C. belong to Achrini, and two others (Melicharella and SIGNALS. Calling signal (Figs 348–354) usually begins Platyproctus) — to Adelungiini. Almost the same groups with a long trill, consisting of hardly distinguishable or merged were outlined in Adelungiinae by Emelyanov [1975], but pulses and sounding like a continuous hiss. Now and then trill is he never gave them any formal rank. I cannot find any interrupted by single high-amplitude pulses or short syllables difference between these two groups in acoustic charac- consisting of 2–5 such ones, each repeating with irregular ters. In representatives of both tribes (or subtribes if intervals (Figs 350, 352). Then male produces a short succes- Adelungiini are regarded as a tribe) calling signals are sion of low-amplitude pulses, delimited both in the beginning and single or repeating phrases (the only exception is M. in the end by several high-amplitude ones, same as in the preceding part of signal (Figs 351, 353). For the most part, song callifrons). Also, in both groups males produce rivalry includes from 1 to 3–4 such successions separated by hissing signals (not registered in P. tessellatus). trills of various length and has overall duration from 10–15 up to Communication systems in Agalliinae and Ade- 30–40 s. lungiinae are similar in several features, but still Courtship behaviour is similar with this, described for M. differ at the same level, as in Agalliinae and Macrop- scanicus by Ossiannilsson [1949]. Singing male mounts female sinae for instance. Both in Agalliinae and in Adelungi- and performs rapid movements by all the body now running a inae male calling signals as a rule are rather complex little forward, now back, now sideways left or right on her back. single or repeating phrases. Thus he runs approximately for 0.5–1.5 s. Then male stops for Their temporal structure somewhat changes during a moment, produces a succession of pulses, similar with this in the second part of calling and resumes his “dancing” again (Figs courtship. However, in Agalliinae these changes are 355–358). It is impossible to ascertain, whether male sings more distinct, besides, male produces copulatory signal during “dancing”, because noises resulting from his movements on the last stage of courtship, whereas in Adelungiinae drown all other sounds. This courtship ritual sometimes lasts for (P. tessellatus) signals of this type were not registered. a minute or two. In all cases observed male leaved female and Also, in contrast with Agalliinae, in males of most ceased singing after several unsuccessful copulation attempts. Adelungiinae competition behaviour is well developed. Several males sitting in one cage produce rivalry signals from So, if any male produces calling, the ones sitting nearby time to time. Signals of this type consist of short trills similar with almost immediately answer him by rivalry signals. these in the second part of calling (Figs 346–347). NOTES. The term “dancing” was also used for description Consequently, judging from acoustic characters, of courtship performance in Alebrini and Empoascini (Typhlo- Agalliinae and Adelungiinae are two groups, having cybinae, see Saxena, Kumar, 1984; Tishechkin, 2001). In rather the rank of subfamilies than tribes. At present, representatives of these tribes, courting male moves trotting by available material does not allow to outline any subdi- small pace and trembling by all the body. Such kind of behaviour visions in the latter group. is quite different from this in Megophthalmus and Ulopa, so these two types of “dancing” must not be confused. Subfamily Megophthalminae There are different opinions concerning position and relationships of Ulopinae. Most authors regard this Megophthalmus scanicus (Fallén, 1806) group as one the most primitive subfamilies among Figs 338–345. Cicadellidae. Evans [1947] places this subfamily at the base of his Ulopides stem which includes also Ledri- LOCALITY. Moscow Area, environs of Pirogovo, meadow on bank of Klyaz’ma River. 29–30.VII, 2.VIII.1987. Signals of 6 ## nae, Hecalinae (Hecalini + Dorycephalini + Eupelicini are recorded at the temperature 22–24°C. + Paradorydiini), Aphrodinae (Aphrodini, incl. Dora- SIGNALS. Calling signal is a train of syllables, repeating turini, Anoterostemma and some others + Errhomenel- with a period about 0.25–0.5 s. The end pulse in each syllable lini, etc.), Cicadellinae and Nirvaninae. Among other as a rule has much higher amplitude, than all other ones. tribes he includes in Ulopinae also Megophthalmini. Several males sitting in the same cage usually sing alter- Wagner [1951] places Ulopinae at the base of his nately. No other exhibitions of competition behaviour were Cicadellides group (Aphrodinae sensu Hamilton [1975] observed. Courtship behaviour in this species was studied by + Penthimiinae + Cicadellidae sensu lato). On the Ossiannilsson [1949]. According to his description, courting other hand, Megophthalminae in his scheme are situat- male emits the same signals as a single one, but he sings more ed at the base of another stem, namely, Iassides (Iassi- regularly, so that pauses between phrases are about as long as the phrase itself. During the pauses he performs rapid jerks nae + Macropsinae + Agalliinae + Idiocerinae). and spasmodic movements with the body, for which the term Linnavuori [1972] also shares the opinion that Ulopi- “dancing” was used by the cited author. After mounting nae are the most primitive group among Cicadellidae. He female male can continue his singing and dancing for about a regards their relationship to Megophthalminae to be ob- minute if immediate attempt of copulation appears to be scure, supposing that the latter group “ may be a unsuccessful. derivative from Ulopinae stock from the Mesozoic era, 48 D.Yu. Tishechkin

338

339 340 2 s 339

200 ms 340 200 ms 341

342 343 344 2 s 342

200 ms 343

200 ms 344

kHz 200 ms 7

6

5 4

3 2 1

0 345 200 ms 346

347 2 s 347 200 ms

Figs 338–347. Vibrational signals of Megophthalmus scanicus (Fall.) (338–345) and Utecha trivia (Germ.) (346–347): 338–344 — oscillograms of calling signals of M. scanicus, 345 — same, oscillogram and sonogram, 346–347 — oscillograms of rivalry signals of U. trivia. Faster oscillograms of the parts of signals indicated as “339–340”, “342–344” and “347” are given under the same numbers. Ðèñ. 338–347. Âèáðàöèîííûå ñèãíàëû Megophthalmus scanicus (Fall.). (338–345) è Utecha trivia (Germ.) (346–347): 338–344 — îñöèëëîãðàììû ïðèçûâíûõ ñèãíàëîâ M. scanicus, 345 — òî æå, îñöèëëîãðàììà è ñîíîãðàììà, 346–347 — ñèãíàëû ñîïåðíè÷åñòâà U. trivia. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “339–340”, “342–344” è “347”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. Vibrational communication in leafhoppers from Ulopides group 49

348 2 s 350 351 349 2 s 353 352 350

200 ms 351 200 ms 352

200 ms 353 kHz 200 ms 7 6 5 4 3 2 1 0 354 200 ms 355

357 2 s 356

358 2 s 357

signal male movements signal 200 ms 358 200 ms

Figs 348–358. Vibrational signals of Utecha trivia (Germ.): 348–353 — oscillograms of calling signals, 354 — same, oscillogram and sonogram, 355–358 — oscillograms of courtship signals. Faster oscillograms of the parts of signals indicated as “350–353” and “357–358” are given under the same numbers. Ðèñ. 348–358. Âèáðàöèîííûå ñèãíàëû Utecha trivia (Germ.): 348–353 — îñöèëëîãðàììû ïðèçûâíûõ ñèãíàëîâ, 354 — òî æå, îñöèëëîãðàììà è ñîíîãðàììà, 355–358 — îñöèëëîãðàììû ñèãíàëîâ óõàæèâàíèÿ. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “350– 353” è “357–358”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 50 D.Yu. Tishechkin but has differentiated so much, that it definitely deserves When several males present on the same twig, they can the rank of separate subfamily”. However, he points out sing alternately. In this situation their signals become distinctly on evident relationship of Megophthalminae and Agalli- shorter. Nevertheless, their specific temporal pattern remains inae. almost unchanged. Davis [1975] considers Agalliinae, Adelungiinae and Female reply is a short (about 0.5 s) trill (Figs 363–364). After Megophthalminae as three closely-related subfamilies, hearing reply signals male starts walking along the plant in but he places Ulopinae somewhat apart from them. different directions, searching for female. During walking he Hamilton [1983] regards Ulopinae as a sister-group clicks the wings from time to time and stops occasionally to produce calling (Fig. 363). If female answers his signal, he usually of Aetalionidae + Membracidae. He joins Megoph- produces one or several additional fragments at the end of her thalminae in one tribe with Agalliinae (under the name reply (about 0.8–1 s after the main part of calling signal, Fig. Megophthalmini) and places them together with Mac- 364). When insects meet, male mounts female, performs short ropsini, Idiocerini, Adelungiini and others in his sub- dancing movements on her back, as male of U. trivia do, and family Eurymelinae. It is necessary to note, that tribes starts producing courtship signal. The signal consists of phrases, in his work for the most part correspond to subfamilies including two parts each. In breaks between phrases 1–2 wings of other authors. clicks usually present (Figs 365–367). Occasionally, on the first Rakitov [1998] joins Ulopinae, Agalliinae, Ade- stage of courtship female answers to each phrase of signal as lungiinae, Macropsinae, Idiocerinae, Megophthalmi- well, as to calling (Figs 368–369). Overall length of courtship nae and some other subfamilies in his group Ulopides. signal sometimes averages several minutes. In the last moments On the phylograms based on molecular data [Di- before copulation male changes his song and repeats second etrich et al., 2001] Ulopinae form a group of closely- part of phrase several times without breaks; thus courtship signal changes into copulatory one (Figs 370–372). Immediately related taxa with Adelungiinae, Agalliinae and Me- after that he makes an attempt to join his genitalia with female’s. gophthalminae. Membracidae appear to be a sister- If the attempt appears to be unsuccessful, male usually starts group of their common stem. courtship signal anew. Also, several authors rise Ulopinae to family rank [Ribaut, 1936; Emelyanov, Kirillova, 1987; Centrotus cornutus (Linnaeus, 1758) Hamilton, 1999). Figs 373–386, 388. In acoustic characters Ulopinae seem to be most closely related with Megophthalminae, due to peculiar LOCALITIES. Moscow Area: (1) Serpukhov District, environs courtship behaviour, including male “dancing”. Avail- of Luzhki Village, 3.VI.1987, signals of 1 # are recorded at the able material on these groups (single species from temperature 21°C; (2) Serpukhov District, environs of Pushchino- each subfamily) does not allow to make any definite na-Oke Town, 9.VI.1987, signals of 2 ## are recorded at the temperature 25–26°C; (3) environs of Pirogovo, 27.V.1988, signals conclusion concerning their relationships with Agalli- of 3 ## are recorded at the temperature 20°C. inae, Macropsinae and Idiocerinae at present. Howev- SIGNALS. Calling signal is a long phrase lasting from 5– er, if the suggestion of several authors about relation of 6 s up to 1 minute and more (Figs 373–375). Its main part is a Megophthalminae with Agalliinae is true, than the com- continuous succession of syllables or pulses with variable mon stem of Ulopinae + Megophthalminae must be temporal pattern (Fig. 376). As a rule, pulses repetition derived from the cluster, formed by Macropsinae, Idio- frequency increases and becomes more stable in the end of cerinae, Agalliinae and related subfamilies. Consequently, succession. Then a short trill of simple discrete pulses follows in this case Ulopinae appear to be a derivative of one of (Figs 377–378, 388). small Cicadellidae stems, but not a sister-group of this Courting male comes close to female and starts singing family in a whole. Similar assumption was expressed more regularly. When producing courtship signal, he attaches additional part consisting of partially merged pulses in the end earlier by Rakitov [1998] and Dietrich et al. [2001]. of each phrase of calling (Figs 379–382). He can sing for several minutes in such a manner. Immediately before and Family Membracidae during copulation attempt male emits copulatory signal con- sisting of alternating last two parts of courtship one (Figs 383– Subfamily Centrotinae 384), i.e. male misses the first prolonged trill. When sitting on female’s back, male performs abrupt jerks by all the body in Gargara genistae (Fabricius, 1775) some appointed moments during singing (Fig. 383). Evidently, Figs 359–372, 387. these jerks are modification of “dancing” observed in G. genistae. When seized or touched by hand, male produces call of LOCALITY. Moscow Area, Serpukhov District, environs of distress (Figs 385–386). Luzhki Village, from Genista tinctoria L. and/or Cytisus ruthenicus Fisch. ex Woloszczak: 19.VII.1988, signals of 2 ## are recorded at the temperature 27°C; 14.VII.1990, signals of 1 # are recorded Subfamily Smiliinae at the temperature 22–23°C; 2–3.VIII.1992, signals of 10 ## and 3 $$ are recorded at the temperature 24–25°C. Stictocephala bisonia Kopp, Yonke, 1977 SIGNALS. Calling signal of this species is a short phrase, Figs 389–403. consisting of succession of pulses and monotonous fragment with indistinct structure (Figs 359–362). These two parts of LOCALITY. Rostov Area, Oblivskiy District, environs of signal differ distinctly from each other by frequency spectra Sosnovy (=Oporny) Village on Chir River, from Rosa sp. on the (Fig. 387). Duration of phrase as a rule averages 1.2–1.5 s. roadside. 18–19.VIII.1992. Signals of 4 ## are recorded at the Phrases follow with irregular intervals. temperature 30–32°C. 200 ms

Vibrational communication in leafhoppers from Ulopides group 51

359 362 2 s 360 361 360 200 ms 361

200 ms 362 200 ms 363

wing clicks 364 200 ms$ $ 2 s 364 wing clicks 200 ms 365

266 367 2 s 366

200 ms 367 200 ms 368 $ 369 2 s 369 200 ms 370

371 4 s 372 371

2 s 372 200 ms

Figs 359–372. Oscillograms of vibrational signals of Gargara genistae (F.): 359–362 — calling signals, 363–364 — male calling and female reply, 365–367 — courtship signals, 368–369 — male courtship signal and female reply, 370–371 — courtship and copulatory signals, 372 — copulatory signal. Faster oscillograms of the parts of signals indicated as “360–362”, “364”, “366–367”, “369”, “371” and “372” are given under the same numbers. Ðèñ. 359–372. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâ Gargara genistae (F.): 359–362 — ïðèçûâíûå ñèãíàëû, 363–364 — ïðèçûâ ñàìöà è îòâåò ñàìêè, 365–367 — ñèãíàëû óõàæèâàíèÿ, 368–369 — ñèãíàë óõàæèâàíèÿ ñàìöà è îòâåò ñàìêè, 370–371 — ñèãíàëû óõàæèâàíèÿ è êîïóëÿöèîííûé ñèãíàë, 372 — êîïóëÿöèîííûé ñèãíàë. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “360–362”, “364”, “366–367”, “369”, “371” è “372”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. 52 D.Yu. Tishechkin

373

374 4 s 374

376 2 s 377 375

378 2 s 376

200 ms 377 200 ms 378 200 ms

379

380 4 s 380 2 s 381 382 381

200 ms 382

male movements 200 ms 383 2 s 384 384 200 ms 385

386 2 s 386 200 ms

Figs 373–386. Oscillograms of vibrational signals of Centrotus cornutus (L.): 373–378 — calling signals, 379–382 — courtship signals, 383–384 — copulatory signals, 385–386 — call of distress. Faster oscillograms of the parts of signals indicated as “374”, “376– 378”, “380–382”, “384” and “386” are given under the same numbers. Ðèñ. 373–386. Îñöèëëîãðàììû âèáðàöèîííûõ ñèãíàëîâCentrotus cornutus(L.): 373–378 — ïðèçûâíûå ñèãíàëû, 379–382 — ñèãíàëû óõàæèâàíèÿ, 383–384 — êîïóëÿöèîííûå ñèãíàëû, 385–386 — ñèãíàë ïðîòåñòà. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “374”, “376–378”, “380–382”, “384” è “386”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. Vibrational communication in leafhoppers from Ulopides group 53

200 ms

0 387

388 200 ms

Figs 387–388. 387 — oscillogram and frequency spectra of different parts of calling signal of Gargara genistae (F.), 388 — oscillogram and sonogram of calling signal of Centrotus cornutus (L.). Ðèñ. 387–388. 387 — îñöèëëîãðàììà è ÷àñòîòíûå ñïåêòðû ðàçëè÷íûõ ÷àñòåé ïðèçûâíîãî ñèãíàëà Gargara genistae (F.), 388 — îñöèëëîãðàììà è ñîíîãðàììà ïðèçûâíîãî ñèãíàëà Centrotus cornutus (L.).

SIGNALS. Only calling signals were registered in my ulated fragments lasting for about 5 s, sometimes experiments. These are single or irregularly repeating phrases followed by several discrete pulses. Normally, it is lasting from 6–10 s up to 1–2 minutes (Figs 389–391, 393, 396). Usually phrase begins with a succession of rather variable male, who walks along the plant searching for female, pulses or syllables. Then syllables repetition rate becomes whereas the latter remains stationary. Any special court- higher and more regular (Figs 389–390, 393, 396). Occasionally, ship signals in this species were not registered. male omits initial part of song (Fig. 391). The phrase ends with Vibrational communication in two American spe- 1–2 syllables separated by intervals about 150–300 ms from the cies of Membracidae was described by Hunt [1993, preceding part of signal. As in other Cicadina, the waveform in 1994]. In Spissistilus festinus (Say) belonging to the syllables depends on physical characteristics of substrate on subfamily Smiliinae, as S. bisonia, communication which the insect occurs. For this reason, the shape of syllables during mating behaviour is similar with this in the latter sometimes varies much in different signals (Figs 392, 394–395, species [Hunt, 1993]. Male produces only one type of 397–399). Several males sitting on the same stem sometimes signal both being single and on all stages of courtship. sing in turn, but never produce rivalry signals. Occasionally, male produces signals with different end part, It is a phrase lasting for approximately 8–9 s and which is a prolonged succession of partially merged syllables consisting of two different parts. The first part is a (Figs 400–403). No any apparent changes in male behaviour pure-tone continuous fragment averaging about 7 s in were observed at these moments. length. Its carrier frequency somewhat changes from the beginning to the end of signal. The second part is a Mating behaviour of S. bisonia was studied by succession of discrete syllables having noise frequen- Strübing [1999]. As in most other Cicadina, receptive cy spectrum. Female reply was observed if the distance female responds to male calling by producing reply between the insects was about 56 cm or less. The pause signals. In this species these are continuous non-mod- between male calling and female reply averaged 2 s. As 54 D.Yu. Tishechkin

389

2 s 390 2 s 391 2 s 392 392

200 ms 393

391 395 2 s 394 200 ms

395 200 ms 396

397 398 399 2 s 397

200 ms 398 200 ms 399

200 ms 400

401 402 403 2 s 401

200 ms 402

200 ms 403 200 ms

Figs 389–403. Oscillograms of calling signals of Stictocephala bisonia Kopp, Yonke. Faster oscillograms of the parts of signals indicated as “392”, “394–395”, “397–399” and “401–403” are given under the same numbers. Ðèñ. 389–403. Îñöèëëîãðàììû ïðèçûâíûõ ñèãíàëîâStictocephala bisonia Kopp, Yonke. Ôðàãìåíòû ñèãíàëîâ, ïîìå÷åííûå öèôðàìè “392”, “394–395”, “397–399” è “401–403”, ïðåäñòàâëåíû ïðè áîëüøåé ñêîðîñòè ðàçâåðòêè íà îñöèëëîãðàììàõ ïîä òàêèìè æå íîìåðàìè. Vibrational communication in leafhoppers from Ulopides group 55 in S. bisonia, after hearing female reply male started mon cluster with Membracidae on cladograms present- walking in different directions searching for the mate, ed in Dietrich et al. [2001]; another cluster nearest to but female remained sitting on the same place. it includes among others Idiocerinae, Eurymelinae and, In Enchenopa binotata Say from the subfamily also, Macropsinae on one of two schemes. Membracinae, communication system is far more com- Both temporal pattern and repertoire (a set of func- plex, than in two previous species. Individual male tional types) of signals in Membracidae are not unique emits calling signals spontaneously . Each signal con- among Membracoidea and does not allow to outline any sists of two sections: the first one is an amplitude- and kind of boundary between this family and Cicadellidae. frequency-modulated wave-train, the second one con- In both species of Centrotinae studied male performs sists of 2 to 10 pulses. Before climbing onto the side rapid, abrupt, jerky movements on female’s back during of female prior to copulation male emits a long and courtship. Among other groups such peculiar behav- elaborate courtship signal. It includes several short iour was observed in Ulopinae (U. trivia) and Megoph- repeating phrases followed by rumbling-like vibration, thalminae (M. scanicus, see Ossiannilsson, 1949) only. same as in the end part of each phrase, but lasting for This seems to be the evidence of relationships between about 5 seconds. Each signal ends by single wing flick. these three groups (Membracidae, Ulopinae and Me- Duration of signal in total exceeds 20 s; usually male gophthalminae). Thus, bioacoustic data corroborate phy- sings for several minutes, repeating his signals. logenetic reconstructions of Rakitov [1998] and Di- Immediately before copulation copulatory signal etrich et al. [2001], but disagree with the opinion, that was detected (the author of the paper cited refers it as Membracidae are sister-group of Cicadellidae as a whole. “second courtship song”). Its amplitude was very low and the identity of individual that produced the signal is Finally, I give a summary of taxonomic conclusions not certain. Copulatory signal consists of two alternat- both from the present article and from two my previous ing parts and lasts less than one minute. ones concerning vibrational communication of Ci- Also, there is a description of rather peculiar behav- cadellidae [Tishechkin, 2000c, 2001]. Several authors iour and parent-offspring communication in subsocial (e.g. Evans, 1947; Wagner, 1951) unite all the subfam- species of the genus Umbonia [Cockroft, 1999]. It is ilies of Cicadellidae into 2–4 groups. In this series of quite possible, that similar communication exists in papers I follow the system of Rakitov [1998]. some species of Aetalion (Aetalionidae), because both Vibrational signals of more than 100 species of imago and nymphs in this genus have well developed Iassides are described in Tishechkin [2000c]. Calling timbals [Evans, 1957]. signals of the most part of Aphrodinae s. str. (Aphro- des, Planaphrodes, Anoscopus and Stroggyloceph- The structure of communication system during alus) are similar with these of Deltocephalini s. str. courtship in Centrotinae (G. genistae) and Membraci- (Deltocephalus, Graminella) and Chelidinus from nae (E. binotata, see Hunt, 1994) is similar, but is quite Paralimnini in general scheme of temporal pattern. By different from this in Smiliinae (C. bisonia and S. this character Deltocephalini (maybe, including Paral- festinus, see Strübing, 1999 and Hunt, 1993). The data imnini) appear to be more closely related with Aphrod- available are inconsistent for making any conclusions inae, than with many other tribes of Deltocephalinae. For at present, but it is quite possible, that acoustic charac- this reason I share the opinion of Hamilton [1975, 1983], ters may be used in taxonomy of Membracidae on the that Aphrodinae and Deltocephalinae must be joined in subfamily or tribe level. one subfamily under the valid name Aphrodinae. Membracidae or Membracidae + Aetalionidae usu- Paralimnini are regarded either as a separate tribe ally are regarded as a sister-group of Cicadellidae (see or as a part of Deltocephalini. These taxa cannot be for instance, Dietrich, Deitz, 1993). Hamilton [1983] separated basing on acoustic characters, so bioacous- divides Cicadellidae in two main branches on his phy- tic data confirm the latter supposition. logram and places Membracidae + Aetalionidae on the Balcluthini (Balclutha punctata (Fabricius, 1775)) top of one branch with Ulopinae as their sister-group. do not differ much from Macrostelini s.str., so the Recently he somewhat modified the scheme: Ulopidae joining of these two tribes in Macrostelini seems to be are raised to family status still remaining as a sister- quite correct. Several Macrosteles species have the group of Membracidae + Aetalionidae, but the com- same scheme of calling signal pattern, as Aphrodini, mon stem of all three families in its turn, is regarded as Deltocephalini and Chelidinus from Paralimnini: the a sister-group of Cicadellidae [Hamilton, 1999]. initial fragment being inconstant in structure and al- Quite different opinion concerning the place of most non-species-specific followed by the main part Membracidae among Membracoidea was expressed by being constant, highly specific and elaborate. The pres- Rakitov [1998] and Dietrich et al. [2001]. Basing on ence of such a variable component may be interpreted morphological, molecular and some other characters as a primitive character, because it cannot play any they suppose that Cicadellidae are paraphyletic with substantial role in a process of recognition of conspe- respect to Membracidae + Aetalionidae, i.e. treehop- cific male due to its non-species-specificity. Recep- pers are specialized line deriving from within Cicadel- tive female never answered to this component, but lidae, and are not their sister-group. Ulopinae, Me- showed high percentage of reply reactions to isolated gophthalminae, Agalliinae and Adelungiinae form com- main part in the experiments with a playback of record- 56 D.Yu. Tishechkin ings of full signals and their different parts [Hunt et al., all Cicadellides. It is impossible to outline any kind of 1992]. Moreover, in several species male occasionally well-defined boundary between these subfamilies bas- misses this component during singing. Thus, it is quite ing on acoustic characters. This fact seems to be the possible, that Aphrodini, Deltocephalini, Paralimnini evidence of their close relationships. Similar opinion and Macrostelini form a group of related tribes, deriv- was expressed by Young [1965] and Hamilton [1983] ing close to the base of common stem of Aphrodinae indirectly by transferring of Mileewini and Nirvanini sensu Hamilton [1975]. Macrostelini seems to be most respectively into Typhlocybinae. primitive group among these tribes due to the presence All Typhlocybinae tribes studied may be arranged in of tremulation fragments in signals of several species. several groups according to the scheme of acoustic Opsiini are separate tribe closely related with Go- communication during mating behaviour. Alebrini and niagnathini, judging by acoustic data. As the four previ- Empoascini form one of the groups. Courtship pattern ous ones, Opsiini + Goniagnathini seemingly form the in these tribes is identical and includes male “dancing”, group, deriving near the base of Aphrodinae stem, but it which was not observed in species from other groups. is not clear, whether it is closely related to Aphrodini Forcipatini sensu Hamilton [1998] and Erythroneurini + Deltocephalini + Paralimnini + Macrostelini. fall into another group: as in Alebrini + Empoascini, The studied representatives of Hecalini (Glosso- courting male in representatives of these tribes pro- cratus foveolatus Fieber, 1866, Hecalus glaucescens duces alternating calling and some additional compo- (Fieber, 1866)), Eupelicini (Eupelix cuspidata (Fabr- nent, but this component is far more developed and icius, 1775)) and Dorycephalini (Dorycephalus hun- united with calling forming an integral signal; more- norum Emeljanov, 1964) are quite similar in temporal over, male does not “dance”. In Vilbasteana oculata pattern of calling signals. Apparently, these taxa must (Lindberg, 1929) (the only studied species of Dikra- be regarded as three closely related tribes, or even as neurini s.str., i.e., excluding Notus and Forcipata) subtribes of a single tribe. Chiasmini (=Doraturini) courtship behaviour is exactly the same as in the previ- seem to be a sister-group of their common stem. ous group, but calling suffers more complex transfor- Calling signals of Nephotettix spp. differ distinctly mation when changes into courtship signal. Typhlocy- from these of other Chiasmini [Inoue, 1982]. So, bini differ from all other tribes either by pattern of Nephotettix and, apparently, also related genera joined courtship behaviour (from Alebrini and Empoascini) or in Stirellini by Emelyanov [1966] represent a separate by temporal structure of courtship signal, which is group, which must be regarded as a tribe or maybe a thoroughly unlike calling in contrast with Erythro- subtribe of Chiasmini. neurini, Forcipatini and Dikraneurini s. str. Athysanini s. str., Cicadulini and Platymetopiini differ in acoustic characters from each other and must Acoustic signals of 50 species of Ulopides (Idio- be regarded as separate tribes. Communication systems cerinae, Macropsinae, Agalliinae, Adelungiinae, Me- in Platymetopius and Fieberiella are identical. For gophthalminae and Ulopinae) and of 3 Membracidae this reason I include Fieberiellini into Platymetopiini ones are described in the present article. following Hamilton [1975]. Idiocerinae seem to be the most primitive among Available data on Grypotini (Grypotes puncticollis groups studied due to the presence of tremulation (Herrich-Schäffer, 1834)) and Selenocephalini (Sele- components in their signals. nocephalus obsoletus (Germar, 1817)) do not allow Macropsinae are most closely related with Agallii- to make any conclusion concerning their status. nae and Adelungiinae. Two latter groups are related, but Iassinae and Penthimiinae are similar with Aphro- separate subfamilies, judging by acoustic characters. dinae sensu Hamilton [1975] in acoustic characters, Distinctive courtship behaviour, which is identical in but, on the other hand, differ from Macropsinae, males of Megophthalminae and Ulopinae, points out on Agalliinae and related subfamilies. So, I place these close relationships between these groups. Similar court- two subfamilies together with Aphrodinae into Ias- ship ritual was also observed in two species of Mem- sides sensu Rakitov [1998]. bracidae studied. This fact corroborates the opinion of Rakitov [1998] and Dietrich et al. [2001], that Mem- Oscillograms and descriptions of vibrational sig- bracidae derives from within Cicadellidae, namely, from nals of 36 species of Cicadellides (Cicadellinae s.l. and Ulopides, but are not a sister-group of this family. Typhlocybinae) are presented in Tishechkin [2001]. Cicadellini (Cicadella, Kolla) differ from all other ACKNOWLEDGEMENTS. I am greatly indebted to Prof. Cicadellinae studied by tremulation mechanism of call- Alexander F. Emelyanov (Zoological Institute of Russian Acad- ing signals producing. For this reason the tribe seems emy of Sciences, St.-Petersburg) for a number of valuable to be the most primitive in the subfamily and standing advises and consultations during the course of this study and for somewhat apart from Errhomenini, Evacanthini and critical reading of the manuscript. On different stages this work was supported by grants of Russian Ministry of Higher Educa- Mileewini. tion (“Investigations in the field of fundamental natural Scienc- Most of Cicadellinae and Typhlocybinae produce es”, head organization — St.-Petersburg State University and pure-tone signals or ones including particular frag- “Universities of Russia”, head organization — Moscow State ments with almost regular sine carrier. Evidently, this University), and of Russian Foundation for Basic Researches character is a synapomorphy of these groups or even of (RFBR, grant No 01–04–49280). Vibrational communication in leafhoppers from Ulopides group 57

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